BERKELEY 

LIBRARY 

UNIVERSITY    Of 

CALIFORNIA 


PALEONTOLOGY  UBRAPV 


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ALASKA 


VOLUME  XIV 


P 


ACH 


841 


ji:y. 


SMITHSONIAN  INSTITUTION 


HARRIMAN  ALASKA  SERIES 

VOLUME    XIV 

Monograph  of  the  Shallow-water 

Starfishes  of  the  North  Pacific  Coast 

from  the  Arctic  Ocean  to  California 

(WITH  110  PLATES) 


ADDISON  EMERY  ^VERRILL 


BY 
VLEl 

Professor  Emeritus  of  Yale  University 

PART  1.    TEXT 


(PUBLICATION  2140) 
CITY  OF  WASHINGTON 

PUBLISHED  BY  THE  SMITHSONIAN  INSTITUTION 
1914 


ttARTH 

SC/ENCES 


Q^afttmore  (p 

BALTIMORE,  MI)..  U.  S.  A 


QL3Z3 
.1 


EDITOR'S  PREFACE. 


The  Starfishes  collected  by  the  Harriman  Alaska  Expedition  were 
sent  to  Professor  A.  E.  Verrill  for  study  and  report  ;  and  through  the 
courtesy  of  various  museums  of  the  United  States  and  Canada 
numerous  additional  specimens  from  the  West  Coast  were  placed  in 
his  hands.  With  this  rich  material  at  his  disposal,  Professor  Verrill 
undertook  a  monographic  revision  of  the  Starfishes  inhabiting  the 
shallow  waters  of  the  Pacific  from  Arctic  Alaska  south  to  California. 

In  the  course  of  three  or  four  years  (by  1904)  the  larger  part  of 
the  monograph  had  been  written,  most  of  the  species  (including 
Stimpson's  types,  in  the  United  States  National  Museum)  had  been 
photographed,  and  more  than  40  plates  had  been  reproduced  and 
printed.  But  pressure  of  other  duties  so  delayed  the  completion  of 
the  work  that  a  dozen  years  have  elapsed  since  it  was  begun.  The 
long  delay,  however,  has  not  been  without  compensations,  for  much 
new  material  has  come  to  Professor  Verrill's  hands,  including  col- 
lections from  Patagonia  and  Fuegia,  enabling  him  to  add  matter  of 
great  importance  on  the  geographic  distribution  of  the  group. 

In  the  meantime,  Mr.  E.  H.  Harriman,  organizer  and  patron  of  the 
Expedition,  having  passed  away,  the  publication  of  the  work,  with 
its  large  series  of  plates,  has  been  generously  provided  for  by  Mrs. 
Harriman. 

In  view  of  Professor  Verrill's  lifelong  studies  of  the  Starfishes,  it  is 
believed  that  the  present  monograph  will  long  remain  a  standard 
authority  on  the  subject,  and  that  it  will  receive  a  hearty  welcome 
from  all  workers  on  the  Echinoderms. 

C.  HART  MERRIAM, 
Editor. 

WASHINGTON,  D.  C, 
May,  1913. 


M886283 


SHALLOW- WATER  STARFISHES  OF  THE 

NORTH  PACIFIC  COAST  FROM  THE 

ARCTIC  OCEAN  TO  CALIFORNIA 


TABLE  OF  CONTENTS. 

PAGE 

Preface    xi 

Introduction    i 

Richness  of  the  Starfish  Fauna ;  Causes I 

Food  and  Feeding  Habits  of  the  Starfishes 3 

Activities  of  Starfishes ;  Migrations ;  Rate  of  Travel 4 

Larval  Stages ;  Protection  of  Eggs  and  Young ;  Incubation 7 

Senses ;  Instincts ;  Memory ;  Acquired  Habits 9 

Peculiarities  of  the  Starfish  Fauna  of  the  Northwest  Coast  of  America  1 1 

Multiplicity  of  Rays ;  Variability  12 

Subspecies  and  Varieties   17 

Hybridism  of  Associated  Species  18 

Faunal  Relations  and  Distribution  of  the  Species 18 

Morphological  Features  of  the  Class  Asterioidea r  20 

Classification  of  the  Asterioidea 24 

Order  Forcipulosa  or  Forcipulata 24 

Various  Kinds  of  Pedicellariae  25 

Families  and  Subfamilies  of  Forcipulosa  26 

Family  Asteriidae  27 

I.  Morphology  and  Growth  of  the  Skeletal  Ossicles  (Text-fig,  i) 27 

II.  Pedicellariae  of  the  Asteriidae 30 

III.  Relative  Importance  of  Morphological  Characters  in  Asteriidae 31 

A.  Modes  of  Growth  and  Increase  in  Number  of  Skeletal  Plates  32 

B.  Interactinal  Plates  and  Spines  33 

C.  Form  and  Arrangement  of  the  Dorsal  Ossicles 34 

D.  Dorsal  Spines ;  Form  and  Arrangement  34 

E.  Pedicellariae ;  Special  Forms  (Text-fig.  2)  35 

F.  The  Madreporite  or  Madreporic  Plate  and  surrounding  Spines  36 

G.  Jaws  and  Oral  Spines  (Text-fig.  3)  36 

H.  Adambulacral   Spines ;   Arrangement    38 

I.  Number  of  Rays ;  Variability 38 

J.  Arrangement  of  the  Ambulacral  Feet  or  Sucker-tubes  (Podia)  39 
K.  Modes  of  Development;  Brooding  of  the  Young;  Position  of 

the  Genital  Pores 39 

Classification   of  Asteriidae    39 

Subfamily  Heliasterinae  40 

Subfamily    Pycnopodiinae    40 

Subfamily  Stichasterinae   40 

Subfamily  Asteriinae  42 

A.  Generic  Subdivisions  of  Asteriinae 43 

B.  Subdivisions  of  the  Asteriinae  having  Definite  Longitudinal 

Rows  of  Dorsal  Plates  and  Spines 44 

C.  Table  of  Extralimital  Species  of  Coscinasterias  and  closely 

Allied  Forms  46 

D.  Observations  on  Various  Genera 46 

(ix) 


TABLE  OF   CONTENTS 


E.  Table  of  Principal  Genera  of  Asteriinae  with  Regular  Longitu- 

dinal Rows  of  Dorsal  Plates  49 

F.  Subdivisions  of  Asteriinae  with  Reticulated  Dorsal  Skeletons. .     50 

G.  Analytical  Table  of  the  Generic  and  Subgeneric  Groups  of 

Northern  Asteriinae  having  Reticulated  Dorsal  Skeletons    54 

H.  Asteriidae  of  the  Northwestern  Coast  of  America 55 

I.  Analytical  Table  of  Genera,  Subgenera,  Species,  and  Subspecies 

of  Asteriidae  from  the  Northwest  Coast  of  America 56 

Asteriinae :     Descriptions  of  Genera  and  Species 67 

Subfamily   Pycnopodiinae    197 

Descriptions    of    Pycnopodiinae 197 

Descriptions  of  Pedicellasteridae 202 

Order  Spinulosa 203 

Morphology    203 

Classification 204 

Suborders  Avelata  and  Velata 204 

Families  of  Spinulosa   204 

Descriptions  of  Families,  Genera,  and  Species 205 

Order   Phanerozona    280 

Morphology   280 

Classification    282 

List  of  Families  282 

Suborder  Valvulosa  or  Valvata  284 

Descriptions  of  Families,  Genera,  and  Species 285 

Suborder  Paxillosa  313 

Descriptions  of  Families,  Genera,  and  Species 314 

Species  erroneously  attributed  to  the  Fauna  337 

Geographical  Distribution 337 

I.  The  Beringian  Fauna  338 

II.  The  Columbia- Alaskan  Fauna 340 

III.  The  Californian  Fauna  343 

IV.  The  South  Californian  Fauna 345 

V.  Some  Comparisons  with  Other  Faunas 347 

VI.  Relations  of  the  Columbia- Alaskan  and  Californian  Faunae 348 

VII.  Relations  of  the  South  Californian  Fauna 349 

VIII.  Patagonian,    Fuegian    and    Antarctic    Shallow-water    Starfishes 

contrasted  with  Those  of  the  North  Pacific  Coast 351 

IX.  List  of  Patagonian  and  Fuegian  Starfishes  Found  in  50  to  500 

Fathoms ;  mostly  Related  to  Northern  or  Arctic  Species 367 

X.  Lists  of   Extralimital   Species   Partially  Described,   Revised,   or 

Figured   370 

1.  North  Atlantic  and  Arctic  Species 370 

2.  Patagonian,  Fuegian,  Antarctic,  and  South  Atlantic  Species  371 

3.  Species  from  the  Panamic  Fauna 372 

4.  Indo-Pacific,  Australian,  and  Japanese  Species 373 

XL  List  of  New  Genera  Established 374 

Bibliography    374 

Index   389 


PREFACE. 

THIS  report  was  originally  planned  to  include  only  the  littoral  and 
shallow-water  species  of  the  starfishes  of  Alaska  and  adjacent  waters 
contained  in  the  fine  collections  made  on  the  Harriman  Expedition  by 
Dr.  (now  Professor)  W.  R.  Coe,  together  with  the  collections  in  the 
Yale  Museum,  previously  received  from  the  same  region,  including 
a  number  of  my  original  types.  The  subsequent  reception  of  numer- 
ous other  large  collections  from  the  same  and  adjacent  regions  soon 
led  to  a  great  extension  of  the  work,  so  as  to  include  all  the  shallow- 
water  species  from  California  to  Bering  Sea.  The  various  collec- 
tions sent  to  me  by  the  Canadian  Geological  Survey  through  the  late 
Mr.  J.  F.  Whiteaves,  and  later  through  Professor  John  Macoun  have 
been  particularly  large  and  valuable,  containing  many  new  forms. 
My  thanks  are  especially  due  to  Professor  Macoun  and  other  mem- 
bers of  the  Canadian  Survey  for  these  favors. 

A  small  but  very  important  collection  was  sent  to  me  from  the 
Museum  of  Comparative  Zoology,  through  the  kindness  of  the 
Director,  Mr.  Samuel  Henshaw,  and  of  Dr.  H.  L.  Clark;  I  also 
enjoyed  an  opportunity  to  study  there  the  entire  collection  of  star- 
fishes from  the  Northwest  coast. 

A  large  collection  was  also  received,  through  Dr.  C.  F.  Newcombe, 
from  the  Provincial  Museum  of  British  Columbia  at  Victoria. 
Professor  Trevor  Kincaid,  University  of  Washington,  Seattle ;  Pro- 
fessor W.  E.  Ritter,  University  of  California,  Berkeley,  and  others 
have  also  sent  useful  collections ;  and  to  all  these  I  wish  to  express 
my  obligations  for  assistance  given. 

A  small  collection,  mostly  from  Bering  Sea,  was  sent  to  me  by 
the  U.  S.  National  Museum,  and  was  very  useful  for  the  more  Arctic 
species.  I  am  also  indebted  to  the  National  Museum,  through  the 
kindness  of  Dr.  R.  Rathbun  and  Miss  M.  J.  Rathbun,  for  photo- 
graphs of  the  type  specimens  of  the  several  species  early  described  by 
Dr.  Wm.  Stimpson,  which  have  been  of  great  use  and  are  now 
reproduced  on  my  plates.  Several  of  these  have  not  been  previously 
figured. 

The  report  was  very  nearly  finished,  mostly  in  its  present  form, 
previous  to  1904,  and  forty-three  plates  had  been  engraved  and 
printed  at  that  time. 

(xi) 


Xll  PREFACE 

In  the  meantime,  Prof.  W.  K.  Fisher,  of  Stanford  University, 
has  published  (1911  b)  an  extensive  work  on  some  of  the  orders  of 
starfishes  of  the  North  Pacific,  including  not  only  very  many  deep- 
water  species,  but  also  considerable  numbers  from  the  shallow  waters 
of  the  same  regions,  covered  by  my  report,  and  including  many  of 
the  species  previously  described  and  figured  by  me,  in  this  work. 

He  had  the  advantage  of  having  studied  my  specimens,  manu- 
scripts and  plates  several  years  ago,  while  visiting  me  at  the  Yale 
Museum.  Yet  in  some  cases  our  conclusions  are  somewhat 
divergent. 

I  have  not  seen  his  types,  nor  have  I  been  able  to  devote  so  much 
time  as  I  would  like  to  the  correlation  of  his  species  of  certain 
difficult  genera,  like  Solaster  and  Henricia,  with  my  own.  But  I  trust 
that  the  figures  and  descriptions  given  in  both  works  will  enable 
future  students  to  understand  what  the  forms  discussed  really  are, 
regardless  of  the  names. 

For  the  numerous  excellent  drawings  of  miscroscopic  details  and 
for  nearly  all  the  photographs,  I  am  indebted  to  the  careful  work  of 
my  son,  Mr.  A.  Hyatt  Verrill,  who  has  devoted  much  time  to  the 
work  during  several  years. 

ADDISON  EMERY  VERRILL. 

NEW  HAVEN,  CONN.,  July,  1912. 


Monograph  of  the  Shallow-water  Starfishes  of  the 

North  Pacific  Coast  from  the  Arctic  Ocean  to 

California,  with  Revisions  of  Various  Extra- 

limital  Genera  and  Species. 


ADDISON  EMERY  VERRILL, 
Professor  Emeritus  of  Yale  University. 


INTRODUCTION. 
RICHNESS  OF  THE  STARFISH  FAUNA;  CAUSES. 

THE  Northwestern  Coast  of  North  America  seems  to  be  the  head- 
quarters or  metropolis  of  shallow-water  starfishes.  No  other  region, 
so  far  as  known,  has  anything  near  so  many  species,  nor  so  many 
generic  and  family  types,  in  shallow  water,  nor  so  great  an  abun- 
dance of  individuals. 

The  species  and  genera  confined  to  the  deeper  waters  off  these 
shores  seem  to  be  equally  numerous,  or  more  so. 

At  present  there  are  over  one  hundred  shallow-water  species 
known  from  that  region,  besides  about  twenty  named  varieties. 

The  abundance  and  variety  of  starfishes  in  these  shallow  waters 
is  apparently  greater  than  at  Panama,  the  West  Indies,  or  the  most 
favorable  parts  of  the  East  Indies  in  similar  depths. 

This  profusion  of  starfishes  does  not  hold  good  for  the  other 
classes  of  Echinoderms.  The  Sea-urchins  (Echinoidea)  are  repre- 
sented by  comparatively  few  species,  of  which  only  about  five  are 
common,  and  those  belong  to  widely  distributed  genera.  But  this 
is  about  the  normal  number  for  any  similar  cold-water  fauna. 

The  serpent  stars  (Ophiuroidea)  are  represented  by  about  the 
usual  number  of  species,  none  of  them  of  unusual  types. 

The  Crinoidea  are  represented  in  shallow  water  only  by  a  single 
large,  comatulid  species,  so  far  as  known  to  me. 

The  Holothurians  are  numerous,  but  not  remarkably  so,  and 
mostly  of  common  genera. 

This  coast  seems  admirably  adapted  for  the  permanent  occupation 
and  evolution  of  starfishes,  and  similarly  for  certain  groups  of  other 
marine  invertebrates  and  fishes. 

2  (i) 


2  VERRILL 

Among  the  fishes  the  Salmonidae  may  be  mentioned.  Among  the 
mollusca  some  groups,  like  the  limpets,  chitons,  Buccinidae,  and 
other  families,  are  much  more  abundant  than  in  any  other  regions. 

Nemerteans  and  tubicolous  Annelids  are  also  remarkably  large  and 
numerous. 

There  is  every  reason  for  thinking  that  such  groups,  including 
various  genera  and  families  of  starfishes,  have  lived  and  flourished 
on  that  coast  for  many  geologic  periods,  and  have  developed  in  that 
region,  by  continuous  evolution,  most  of  the  species  found  there  at 
present,  as  well  as  great  numbers  that  have  disappeared. 

Indeed,  I  am  led  to  conclude  that  this  region  has  been,  for  long 
ages,  a  center  of  evolution,  from  which  various  generic  and  specific 
types  of  starfishes  have  been  dispersed  to  other  faunal  areas,  with 
or  without  subsequent  changes. 

It  is  not  at  all  improbable  that  some  arctic  and  boreal  species  of 
the  North  Atlantic  originated  on  the  Alaskan  coast  and  have  since 
crossed  to  the  Atlantic  by  way  of  the  Arctic  Ocean,  when  the  arctic 
climate  was  warmer. 

Others  may  have  migrated  southward  to  the  subtropical  and 
tropical  regions  of  Mexico  and  Central  America,  and  even  to  the 
Antarctic  Seas,  for  some  of  the  deep-water  Patagonian  and  antarctic 
species  show  peculiar  affinities  with  the  Alaskan  species. 

Such  migrations,  across  the  tropics,  would  easily  be  possible  for 
species  having  a  considerable  range  in  depth,  for  their  congenial 
temperatures  could  easily  be  found  at  no  great  depth,  even  off 
Panama. 

One  reason  for  the  great  variety  and  persistence  of  the  starfishes 
and  other  groups  on  this  coast  is,  no  doubt,  the  fact  that  the  tem- 
peratures of  the  waters  were  not  seriously  reduced  during  the 
Glacial  Period,  while  those  of  the  North  Atlantic  were  rendered  so 
cold  that  the  preexisting  fauna  was  largely  exterminated,  while  the 
survivors  were  driven  southward,  except  in  the  case  of  eminently 
arctic  and  deep-sea  species. 

Another  very  favorable  condition  is  the  comparatively  equable 
temperature  of  those  waters,  due  to  the  reflex  of  the  great  Japanese 
tropical  current  of  water.  Its  influence  in  modifying  the  climate 
of  the  whole  coast,  south  of  the  Aleutian  Islands,  at  least,  is  well 
known. 

For  the  prosperity  of  most  marine  animals,  equability  of  temper- 
ature is  exceedingly  favorable,  whether  high  or  low,  especially  at  the 
breeding  season. 


SHALLOW-WATER   STARFISHES  3 

No  doubt  the  comparative  uniformity  of  the  temperatures  at  all 
seasons,  and  over  vast  areas,  in  the  deep  sea,  is  the  principal  cause 
of  the  great  abundance  of  starfishes  and  many  other  groups  of 
animals  in  the  deep  sea  and  of  their  wide  distribution. 

Constancy  of  temperature  at  the  breeding  season,  year  after  year, 
is  of  the  greatest  importance  to  nearly  all  animals,  both  in  the  sea 
and  out  of  it,  as  the  writer  pointed  out  many  years  ago  in  the  case 
of  birds,*  and  as  has  since  been  confirmed  by  other  investigators  for 
birds  and  other  groups. 

Another  very  favorable  condition,  for  the  multiplication  of  star- 
fishes on  this  coast,  is  the  vast  extent  and  broken  condition  and  rocky 
character  of  the  coast-line,  with  innumerable  islands,  bays,  straits, 
fiords,  and  inlets,  affording  every  variety  of  stations  and  any  amount 
of  shelter  from  severe  storms,  and  at  the  same  time  furnishing 
innumerable  suitable  stations  on  the  rocky  shores,  for  the  growth 
and  increase  of  all  sorts  of  plant  and  animal  life,  on  which  the  star- 
fishes may  feed.  It  has  been  stated  that  the  coast-line  of  Alaska  and 
the  adjacent  islands  exceeds  26,000  miles,  or  more  than  the  circum- 
ference of  the  earth.  That  of  British  Columbia  is  also  of  vast  extent. 

FOOD  AND  FEEDING  HABITS  OF  THE  STARFISHES. 

The  littoral  and  shallow-water  starfishes  are  nearly  all  carnivorous, 
by  preference,  and  feed  very  largely  on  the  barnacles  and  mollusca 
that  live  among,  or  attached  to,  the  rocks,  such  as  mussels,  oysters, 
limpets,  chitons,  small  spiral  gastropods,  etc. 

Considering  the  great  numbers  and  large  sizes  of  many  of  the 
starfishes,  the  wonder  is  that  they  have  not  already  entirely  exter- 
minated those  mollusca  on  which  they  largely  feed. 

The  vast  amount  of  damage  done  to  the  cultivated  oyster  beds  on 
our  Atlantic  coast  by  a  single  small  species  of  starfish  (Asterias 
forbesi)  is  well  known.  What  then  must  be  the  destruction  wrought 
to  the  bivalves  on  the  Pacific  coast,  where  there  are  some  forty 
related  species,  with  similar  habits,  many  of  them  becoming  over 
two  feet  across  ? 

One  of  our  native  starfishes,  six  inches  across,  will  eat  over  twenty 
small  oysters  in  one  day.  Probably  it  would  take  a  hundred  oysters 
or  mussels  to  satisfy  one  of  the  giant  starfishes  of  Alaska  or  Puget 
Sound. 

1  Proceedings  Boston  Soc.  Nat.  Hist.,  vol.  x,  p.  259,  1866. 


4  VERRILL 

The  great  abundance,  size,  and  voracity  of  these  Pacific  starfishes 
will  always  be  a  great  hindrance  to  the  cultivation  of  oysters,  mus- 
sels, etc.,  on  that  coast,  except,  perhaps,  where  the  water  is  dis- 
tinctly too  brackish  for  starfish  life. 

Fortunately  for  the  mollusca,  at  least,  most  of  the  larger  littoral 
starfishes  are  more  or  less  cannibals.  When  opportunity  occurs, 
they  do  not  hesitate  to  devour  each  other. 

From  the  stomach  of  a  large  ten-armed  Solaster,  I  have  taken  the 
half-digested  ray  of  an  Asterias  that  must  have  been  a  foot  across 
when  living.  An  inch  or  two  of  the  ray  still  protruded  from  the 
mouth  of  the  Solaster,  for  it  was  much  too  large  to  swallow  entire. 

The  various  kinds  of  starfishes  that  inhabit  rocky  shores,  clinging 
to  the  rocks,  have  large  numbers  of  strong,  muscular  sucker-feet, 
each  tipped  with  a  perfect  sucker  for  adhesion.  In  the  ordinary 
five-rayed  kinds,  six  to  eight  inches  across,  there  will  be  four  close 
rows  of  such  suckers  the  whole  length  of  the  ray,  perhaps  200  to  a 
ray;  or  1,000  on  the  five  rays. 

In  the  case  of  the  many  larger  five-rayed  and  six-rayed  kinds,  two 
feet  across,  the  number  of  suckers  may  be  4,000  to  8,000  or  more. 

How  many  there  are  on  the  big  twenty-rayed  or  twenty-four- 
rayed  star  of  that  coast,  which  becomes  over  thirty  inches  across,  and 
has  the  suckers  mostly  crowded  into  four  rows,  on  each  ray,  nobody 
knows,  for  apparently  nobody  has  had  the  patience  to  count  them. 
Probably  there  may  be  more  than  40,000.  I  refer  to  the  great  Pycno- 
podia  or  "  sun-star."  (See  pi.  xxix.) 

Such  starfishes  are  able  to  open  an  oyster  or  mussel  by  a  long  and 
steady  pull  with  these  suckers,  while  the  rays  are  wrapped  around 
the  victim.  All  such  starfishes  can  evert  the  large,  loose,  bag-like 
stomach  and  wrap  it  around  its  prey  till  digestion  is  completed, 
if  it  be  too  large  to  swallow  entire.  But  the  mouth  is  also 
very  extensible  and  dilatable,  so  that  they  can  swallow  objects  sur- 
prisingly large.  I  have  often  taken  sea-urchins  an  inch  or  more  in 
diameter  with  the  spines  nearly  all  in  place,  from  the  stomachs  of 
starfishes  of  no  more  than  ordinary  size. 

ACTIVITIES  OF  STARFISHES;  MIGRATIONS;  RATE  OF  TRAVEL. 

Although  most  starfishes,  as  ordinarily  seen  in  life  clinging  to 
stones,  etc.,  appear  very  sluggish  and  slow  in  their  motions,  they 
are  really  able  to  travel  to  considerable  distances  and  undoubtedly, 
in  some  cases,  make  migrations  of  considerable  extent,  either  in 


SHALLOW-WATER   STARFISHES  5 

search  of  food  or  to  seek  more  congenial  temperatures  in  deeper  or 
shallower  water,  or  for  other  reasons,  such  as  breeding  habits. 

In  Long  Island  Sound  it  is  a  common  experience  of  the  oyster 
cultivators  to  find  that  many  thousands  of  starfishes  (Asterias  for- 
besi)  suddenly  appear  on  certain  oyster  grounds  which  had  been 
nearly  or  quite  free  of  them  a  few  days  previously.  If  undisturbed 
for  a  few  weeks  they  may  destroy  thousands  of  bushels  of  oysters 
and  then  as  suddenly  disappear  from  the  devastated  grounds,  to  visit 
some  other  locality. 

In  the  autumn,  September  to  November,  as  I  have  myself 
observed,  year  after  year,  at  Outer  Island,  of  the  Thimble  Islands 
group,  in  Long  Island  Sound,  they  migrate  upward  from  deeper  to 
shallower  water,  and  may  then  be  found  in  myriads  on  rocky 
shores  between  tides,  where  none  were  to  be  seen  a  few  days  before. 

At  such  times  large  numbers  are  often  caught  and  destroyed  in 
the  interests  of  the  owners  of  oyster  grounds  in  the  vicinity.  In 
some  seasons  a  boy,  one  of  my  grandsons,  often  caught  during  low 
tide  two  or  three  bushels,  morning  after  morning,  on  one  small 
island,  up  to  a  total  of  fifteen  or  sixteen  bushels,  and  more  still  came 
up  every  morning.  These  were  mostly  not  more  than  half-grown 
and  a  bushel  would  usually  count  up  to  1,000  or  more.  The  same 
operation  was  carried  on  at  many  adjacent  islands  by  others,  with 
similar  results.  The  effect  of  this  wholesale  slaughter  was  said  to 
have  notably  decreased  the  loss  of  oysters  on  the  oyster  grounds  of 
the  district  during  the  next  year.1 

At  this  time  the  starfishes  were  feeding  mainly  on  the  young 
oysters  of  the  season,  which  had  "  set "  thickly  on  all  the  rocks  and 
ledges,  below  half-tide,  but  they  were  also,  in  many  cases,  feeding 
on  the  rock  barnacles  (Balanus  balanoides),  which  were  also  abun- 
dant on  the  rocks. 

Later  in  the  season  they  disappear,  going  into  deeper  water  to 
avoid  freezing  temperatures. 

I  have  repeatedly  tested  their  rate  of  travel,  on  the  shores  of 
Outer  Island.  By  placing  a  hundred  or  more  in  a  small  pool  of 
water  left  by  the  tide  at  about  half-tide  mark,  it  was  easy  to 
ascertain  how  far  and  in  what  directions  they  would  travel  during 

1  Immense  numbers  of  this  same  starfish  are  also  taken  in  Long  Island 
Sound  and  other  oyster-growing  waters  by  dragging  "  tangles  "  or  "  swabs  " 
over  the  oyster  beds,  as  first  recommended  by  the  writer  in  1873  and  1876. 
A  single  oyster  steamer  by  this  means  has  often  taken  twenty-five  barrels 
or  more  in  a  short  time. 


O  VERRILL 

about  six  hours,  while  the  tide  was  above  the  outlet  of  the  pool, 
which  was  on  a  nearly  smooth,  gently  sloping,  granite  ledge. 

In  brief,  nearly  all  left  the  pool,  and  travelled  downward,  but  in 
irregularly  divergent  lines,  to  various  distances,  varying  from  one 
to  about  fifteen  feet.  The  majority  followed  a  nearly  straight 
course  down  the  more  sloping  parts  of  the  ledge,  but  were  readily 
diverted  to  one  side  or  the  other  by  small  variations  in  the  slope. 
Those  that  travelled  the  greater  distances  probably  left  the  pool 
soonest,  and  were  perhaps  in  more  active  condition.  Selecting  a 
dozen  or  so  of  the  best  travellers,  they  might  average  ten  feet  to 
twelve  feet  in  six  hours,  or  nearly  two  feet  an  hour,  providing  th§y 
left  the  pool  about  as  soon  as  it  was  covered  by  the  tide,  which  was 
probably  true  of  some  of  them,  at  least.  As  their  motions  are 
rather  deliberate  and  continuous,  they  could,  no  doubt,  keep  up  this 
rate  for  many  hours,  or  about  336  feet  in  a  week. 

Yet  these  were  mostly  rather  small,  from  4  to  6  inches  across,  and 
the  larger  ones  were  found  to  travel  much  faster  than  the  young 
ones,  for  they  have  a  much  larger  number  of  sucker-feet,  which  are 
also  stronger  and  longer. 

Such  observations  as  the  above  give,  of  course,  only  an  approxi- 
mate idea  of  their  rate  of  travel.  These  were  neither  impelled  by 
hunger  nor  by  cold,  but  merely  by  an  instinctive  tendency  to  seek  a 
lower  level.  A  stronger  stimulus  might  have  increased  the  speed. 

Very  large  starfishes,  like  many  of  the  west  coast  species,  would 
probably  travel  very  much  faster,  and  perhaps  those  with  many  rays 
and  a  vast  number  of  suckers,  like  Pycnopodia  and  Solaster,  would 
be  still  more  speedy.  By  actual  count  a  half-grown  Pycnopodia  was 
found  by  the  writer  to  have  over  22,000  ambulacral  feet. 

At  any  rate,  the  observed  speed  would  be  quite  sufficient  to  permit 
them  to  ascend  or  descend  a  sloping  sea  bottom  quickly  enough  to 
avoid  any  ordinary  changes  in  the  temperature  of  the  water  liable  to 
be  injurious,  and  also  sufficient  to  enable  them  to  seek  their  ordinary 
food  on  new  grounds.  It  is  known  that,  in  spite  of  their  voracity, 
they  can  go  for  many  days,  or  even  weeks,  without  any  food  and  still 
remain  active  and  in  good  health. 

However,  there  are  other  families  of  starfishes  that  are  far  more 
lively,  while  some  others  are  far  more  sluggish. 

The  most  lively  starfishes  known  to  me  belong  to  the  genus 
Luidia.  Some  of  these  are  very  large  species. 

These  live  on  bottoms  of  fine  sand  or  mud,  more  commonly  in 
rather  deep  water,  but  often  close  to  the  shore  in  sheltered,  shallow, 


SHALLOW-WATER   STARFISHES  7 

sandy  bays  and  harbors.  Some  of  the  species  have  more  than  six 
rays.  In  all  cases  the  rays  are  long,  flattened,  and  very  flexible. 
Ordinarily  when  at  rest  they  lie  just  beneath  the  surface  of  the  sand, 
which  soon  forms  into  star-like  impressions,  agreeing  with  the  star- 
fish in  size  and  form.  This  is  due  to  the  minute  currents  of  water 
produced  by  the  ciliary  motions  connected  with  the  respiration  of 
the  starfish  by  means  of  the  dorsal  papulae. 

When  disturbed,  the  starfish  glides  away  quickly,  just  under  the 
loose  superficial  sand,  using  its  "  sucker-feet  "  or  "  ambulacral  feet " 
as  paddles  for  swimming  or  gliding.  These  ambulacral  feet  are 
much  larger  and  longer  than  usual  and  are  much  flattened  trans- 
versely, and  have  no  terminal  sucker,  but  end  in  a  point.  They  are 
very  muscular  and  can  be  waved  back  and  forth,  like  paddles,  in 
unison,  so  that  the  motion  is  more  like  rowing  or  swimming  than 
running  or  creeping.  While  under  the  sand,  no  doubt  the  pressure 
against  the  sand  causes  them  to  act  something  like  legs  or  feet ;  but 
when  placed  in  an  aquarium  without  sand,  they  can  swim  or  glide 
along  the  bottom  and  up  the  perpendicular  sides  with  surprising 
rapidity. 

My  own  observations  on  this  genus  were  made  in  1901,  at  Ber- 
muda *  on  Luidia  clathrata  Say,  a  five-rayed  species. 

Similar  observations  have  since  then  been  made  by  others  on 
different  species  of  Luidia,  so  that  this  mode  of  locomotion  is  prob- 
ably common  to  all  the  species  of  the  genus. 

LARVAL  STAGES;  PROTECTION  OF  EGGS  AND  YOUNG; 
INCUBATION. 

The  wide  and  rapid  distribution  of  many,  if  not  most,  starfishes  of 
shallow  seas  is,  however,  due  to  the  fact  that  nearly  all  produce  vast 
numbers  of  minute  eggs,  which  develop  into  small,  singular,  bilateral, 
free-swimming  larval  forms  (bipinnaria,  brachiolaria)  that  require 
many  days  or  weeks  to  go  through  their  metamorphoses  before  set- 
tling down  to  the  bottom  in  the  starfish  form.  While  in  these  larval 
stages,  they  may  be  drifted  long  distances  by  waves  and  currents  and 
finally  settle  down  in  places  far  from  their  place  of  origin.  Of  course, 
in  most  cases  they  become  widely  dispersed  and  vast  numbers  perish, 
but  sometimes,  on  the  other  hand,  myriads  of  young  may  happen  to 
drift  along  in  company,  by  the  action  of  a  current,  and  so  finally 
locate  as  a  vast  colony  in  a  new  home. 

1  A.  E.  Verrill,  1901,  p.  36,  and  Zoology  of  Bermuda,  Article  10,  p.  36,  1903. 
(See  Bibliography.) 


8  VERRILL 

Perhaps  the  sudden  appearance  of  great  numbers  of  young  star- 
fishes on  certain  oyster  grounds,  where  they  were  not  found  pre- 
viously, may  be  accounted  for  in  this  way.  When  well  fed  on  young 
oysters,  these  little  baby  starfishes  grow  rapidly  and  soon  become 
large  enough  to  attract  attention  and  do  great  mischief,  even  in  one 
season.1 

Certain  genera  and  species  of  shallow-water  starfishes,  and  per- 
haps a  large  proportion  of  the  deep-sea  species,  do  not  produce  so 
many  minute  eggs,  nor  do  these  develop  into  free-swimming  larvae. 
On  the  contrary,  the  mother  retains  the  eggs  and  cares  for  them  till 
they  pass  through  an  abbreviated  metamorphosis  and  develop  into 
minute  young  starfishes  provided  with  "  sucker-feet "  or  podia  to 
enable  them  to  take  care  of  themselves. 

Commonly,  in  such  cases,  the  eggs  and  young  are  held  under  and 
around  the  mouth  in  large  clusters.  During  the  time  required  for 
their  development  the  mother  appears  to  be  incapable  of  taking  any 
food,  owing  to  the  obstruction  of  the  oral  region.  The  young  are 
often  attached  by  a  larval  actinal  pedicel  in  clusters  or  strings. 

This  is  the  mode  of  carrying  the  eggs  and  young  observed  in  sev- 
eral species  of  Henricia,  Anasterias,  Sporasterias,  and  Podasterias, 
and  in  numerous  small  species  of  Asteriidae,  belonging  to  the  genus 
Leptasterias.  I  have  personally  observed  this  in  L.  compta,  L.  tenera, 
and  L.  tittoralis  of  the  New  England  coast,  and  in  L.  epichlora  and 
cribraria  of  Alaska.  The  young  of  epichlora  were  collected  by  Dr. 
W.  R.  Coe.  (See  pi.  LXXXV,  figs.  2,  a-/.)  In  all  these  the  genital 
pores  are  on  the  ventral  side,  near  the  mouth. 

In  some  other  cases  (Leptychaster)  the  mother  carries  the  eggs  on 
the  back,  between  the  spines. 

Another  method  occurs  in  the  genus  Pteraster,  and  probably  in  all 
the  related  genera  of  the  family  Pterasteridae.  In  these  there  is 
a  large  dorsal  "  nidamental  pouch,"  marsupium,  or  better,  gono- 
codium,  for  the  retention  of  the  eggs  and  the  development  of  the 
young,  which  remain  in  it  until  fully  formed  and  sometimes  up  to 
10  to  15  mm.  or  more  in  diameter. 

This  gonocodium  is  formed  by  a  tent-like  membrane,  which  is 
supported  and  kept  stretched  by  the  tips  of  slender,  elongated, 
radiating  spinules,  arising  from  the  dorsal  ossicles,  and  long  enough 

1  Starfishes  that  have  such  minute  eggs  and  young  usually  have  minute 
genital  pores,  situated  in  pairs,  dorsally,  in  the  interradial  areas.  Those 
that  have  large  eggs  that  they  carry  around  or  over  the  mouth  have  the 
genital  pores  larger,  on  the  ventral  side  near  the  mouth,  so  far  as  observed. 


SHALLOW-WATER   STARFISHES  9 

> 

to  leave  a  considerable  cavity  beneath  it,  in  which  the  genital  orifices 
and  respiratory  papulae  are  situated.  This  membrane  is  perforated 
by  small  pores  for  the  ingress  of  water,  and  normally  has  a  large, 
valvular,  central  oscule  through  which  the  water  escapes.  The  mem- 
brane has  muscular  fibers  for  contraction. 

The  eggs  are  laid  and  fertilized  in  the  gonocodium,  and  when  the 
young  are  matured  they  often,  or  perhaps  always,  escape  by  rup- 
tures of  the  membrane,  which  later  heal  up — a  sort  of  natural  caesa- 
rian  operation. 

In  this  group  the  eggs  are  few  in  number,  as  usual  in  all  cases 
where  the  young  are  well  cared  for.  Several  species  of  this  family 
occur  on  the  west  coast.  (See  pi.  xxxn.) 

SENSES;  INSTINCTS;  MEMORY;  ACQUIRED  HABITS. 

Comparatively  little  is  known,  from  actual  observation,  as  to  the 
nature  or  acuteness  of  the  senses  in  starfishes. 

At  the  tip  of  each  ray,  in  the  apical  plate,  there  is  a  small  and 
very  simple  eye,  or  pigmented  eye-spot,  often  red  in  color  during 
life.  .  It  is  at  the  end  of  the  radial  nerve.  Probably  this  kind  of  an 
eye  is  of  use  only  for  distinguishing  different  degrees  of  light,  or 
possibly  the  shadow  of  a  nearby  moving  object. 

Close  to  the  eye  there  is  a  single  odd  ambulacral  tube,  generally 
considered  a  tentacle,  and  supposed  to  have,  at  least,  the  sense  of 
touch.  However,  all  parts  of  the  integument,  and  especially  the 
papulae  and  sucker-feet,  are  also  very  sensitive  to  touch.  There- 
fore it  may  be  that  these  tentacles  have  some  other  sense  func- 
tion more  developed  than  elsewhere,  possibly  that  analogous  to 
smell  or  taste,  for  appreciating  the  purity  or  salinity  of  the  water. 
They  must  have  a  sense  for  heat  and  cold,  as  this  determines  their 
migrations  at  certain  seasons.  That  they  are  somewhat  sagacious 
and  discriminating  in  their  successful  selection  of  the  young  oysters 
for  food  in  spite  of  the  labor  and  time  required  to  open  them,  is 
good  evidence  that  they  have  some  sense  analogous  to  taste,  and  have 
sense  enough  to  follow  it. 

That  they  can  be  attracted  to  an  open  oyster  or  other  favorite 
food,  from  some  distance,  indicates  that  they  have  some  sense  simi- 
lar to  smell,  like  that  of  many  mollusca  and  Crustacea  that  are  easily 
attracted  by  a  dead  fish  or  other  odorous  food.  As  they  have  no 
proper  head  or  central  brain,  but  only  a  circle  of  equal  nervous 
centers,  it  is  clear  that  they  can  have  no  consciousness  of  right  or 
left,  forward  or  backward.  Each  ray  is  practically  equal  to  every 


IO  VERRILL 

other,  yet  all  the  vast  number  of  suckers,  sometimes  20,000  or  more, 
and  all  the  rays  must  act  in  unison  or  there  could  be  no  progression, 
nor  any  procuring  of  food. 

Just  how  starfishes  decide  to  travel  in  one  direction  rather  than 
in  another  is  a  problem  difficult  to  solve.1 

Dr.  Jennings,  who  has  experimented  extensively  with  reference 
to  the  behavior  of  starfishes,  says  very  truly  that  whatever  a  starfish 
can  do  at  all  it  can  do  in  many  different  ways  and  seldom  does  it 
twice  in  just  the  same  way.  This  is  particularly  applicable  to  its 
modes  of  righting  itself  when  turned  over,  getting  out  from  under 
weights,  elastic  bands,  etc.  He  found  that  when  a  starfish  had  been 
forced  to  use  the  same  arms  to  right  itself  very  many  times  and  for 
several  weeks,  it  could  thus  be  trained  to  continue  to  use  the  same 
arms,  and  thus  had  acquired  a  new  habit,  but  if  left  to  itself  it  lost  the 
habit  in  about  a  week. 

Perhaps  the  preference  of  the  starfish  to  use  the  same  rays  was 
mainly  because,  by  repeated  use,  the  muscles  of  those  rays  had  become 
somewhat  larger  and  stronger  by  the  systematic  "  training." 

I  know  of  no  other  successful  attempts  to  educate  a  starfish.  In 
nature  they  seem  to  show  some  memory  and  some  persistency.  I 
once  placed  a  large  active  holothurian  (Thy one  briar  eus],  that  I 
wished  to  figure,  in  one  end  of  a  large  aquarium,  about  four  feet 
long,  while  there  was  a  starfish  (Asterias  forbesi)  at  the  other  end. 
Next  morning  the  starfish  was  mounted  on  the  Thyone  and  had 
slightly  eroded  its  skin  by  means  of  its  oral  spines.  I  disengaged  the 
Asterias,  put  it  at  the  opposite  end  of  the  tank  with  stones  and  other 
obstacles  between,  and  supplied  other  food,  such  as  cracked  mus- 
sels, etc.,  and  other  Thy  ones,  apparently  just  as  good.  In  a  couple 
of  hours  it  was  back  again,  working  away  at  the  same  spot  on  the 
holothurian.  The  same  course  was  taken  a  second  time,  with  the 
same  result.  The  starfish  was  evidently  bound  to  eat  that  particular 
Thyone  or  go  without  any  dinner.  I  had  to  remove  it  from  the 
tank  to  save  the  Thyone.  Apparently  the  Asterias  either  remem- 
bered where  his  chosen  dinner  rested,  or  else  it  had  a  keen  sense  of 
smell  to  distinguish  it  from  others  of  the  same  species,  and  from 
other  natural  food. 

In  respect  to  maternal  instincts,  the  most  interesting  case  known 
to  me  was  told  to  me  by  Prof.  Louis  Agassiz,  while  I  was  a  student 

'  Dr.  L.  J.  Cole  ( Journ.  Exper.  Zool,  xiv,  No.  I,  1913)  has  recently  made 
some  interesting  experiments  on  the  behavior  of  Asterias  forbesi  under 
special  conditions,  as  to  the  relative  use  of  the  various  arms  as  anterior  or 
directive,  etc. 


SHALLOW-WATER   STARFISHES  II 

and  assistant  with  him  about  1862.  I  am  not  aware  that  it  has  been 
published.  It  is  recorded  in  a  diary  kept  by  me  at  that  time. 

He  said  that  he  had  a  large  aquarium,  at  Nahant,  Mass.,  in  his 
summer  residence,  I  think,  and  placed  in  it  a  large  female  Solaster 
endeca*  which  was  carrying  a  cluster  of  eggs  over  the  oral  area.  As 
an  experiment,  he  removed  the  cluster  and  put  it  at  the  extreme 
opposite  end  of  the  aquarium.  In  a  few  hours  the  starfish  found 
the  cluster  and  replaced  it  as  at  first. 

Mr.  E.  Desor  (Proc.  Boston  Soc.  Nat.  Hist.,  vol.  in,  p.  n,  1848) 
records  a  similar  observation  in  the  case  of  Henricia  sanguinolenta. 

I  have,  personally,  never  had  a  chance  to  repeat  this  experiment, 
for  though  I  have  dredged  thousands  of  Solasters,  it  has  always 
been  at  the  season  of  the  year  when  they  were  not  carrying  eggs. 
Such  experiments  should  be  repeated  at  the  various  permanent 
seaside  laboratories  now  in  existence  on  both  coasts.  None  existed 
during  the  long  period  of  most  of  my  field  work,  except,  after  1881, 
that  of  the  U.  S.  Fish  Commission. 


PECULIARITIES  OF  THE  STARFISH  FAUNA  OF  THE  NORTH- 
WEST COAST  OF  AMERICA. 

The  general  discussion  of  the  geographical  distribution  of  the 
genera  and  species  and  their  relations  to  those  of  other  regions  must 
be  left  to  a  final  chapter.  There  are,  however,  some  very  peculiar 
features  of  more  general  interest,  to  which  it  seems  best  to  call 
attention  in  this  place. 

The  great  size  of  many  of  the  species,  some  of  which  are  among 
the  largest,  if  not  the  very  largest  species  known,  has  already  been 
mentioned.* 

The  Pycnopodia,  though  perhaps  not  the  heaviest,  is  probably 
the  largest  starfish  known.  It  becomes  at  least  thirty-six  inches 
broad ;  and  according  to  some  collectors  may  be  even  four  feet  across. 

*As  no  one  else  has  recorded  the -carrying  of  eggs  by  Solaster,  and  as 
its  genital  pores  are  described  as  minute  and  dorsal,  I  suspect  that  there 
was  an  error  on  my  part  in  recording  the  name,  or  on  his  part  in  giying  it. 
Perhaps  Henricia  was  meant. 

*  At  least  nine  of  the  species  of  Asterias  and  allies  become  there  over  two 
feet  in  diameter.  Among  these  large  species  are  Orthasterias  Columbians 
Ver.,  O.  forcipulata  Ver.,  Ev.  troschelii  (St.),  var.  rudis  V.,  Ev.  acanthostoma 
V.,  Pisaster  ochraceus  (Br.),  P.  giganteus  (St.),  P.  liitkenii  (St.),  P.  papulosus 
Ver.,  Pycnopodia  helianthoides  (Br.).  Luidia  foliolata  becomes  about  as 
large. 


12  VERRILL 

MULTIPLICITY  OF  RAYS ;  VARIABILITY. 

It  is  a  singular  fact,  for  which  no  satisfactory  reason  has  yet  been 
given,  that  nearly  all  Echinoderms,  whether  living  or  fossil,  are  nor- 
mally five-rayed.  This  rule  prevailed  quite  as  constantly  in  the 
remote  paleozoic  ages  as  in  the  present  period. 

The  number  of  rays  was  apparently  well  fixed  in  the  unknown 
primordial  ancestors  of  the  earliest  fossil  Echinoderms  of  most 
classes,  for  all  classes  of  Echinoderms,  except,  perhaps,  the  Holo- 
thurians,  appeared  in  the  early  geologic  ages,  with  most  of  their 
more  important  features  much  as  we  find  them  now. 

Among  Paleozoic  Echinoderms,  as  among  modern  ones,  certain 
starfishes  departed  from  the  five-rayed  type  by  acquiring  additional 
rays.  As  in  some  modern  genera,  this  was  apparently  done  in  post- 
larval  life  by  the  interpolation,  or  budding  in,  of  new  rays  between 
the  older  ones,  in  some  of  those  species  having  numerous  rays. 

At  present,  this  is  known  to  occur  only  in  Pycnopodia  of  the 
Northwest  coast ;  in  Heliaster,  represented  by  several  species  on  the 
tropical  and  subtropical  Pacific  coasts,  from  lower  California  to 
Chile ;  and  in  Labidiaster,  from  Patagonia.  It  is  probable,  also,  that 
it  occurs  in  Rathbunaster  Fisher,  a  deep-water  Californian  species, 
allied  to  Pycnopodia. 

Of  Heliaster,  seven  species  are  known.  When  adult,  the  number 
of  rays  exceeds  twenty  and  in  some  species  is  as  high  as  forty  to 
forty-four. 

In  all  its  species  new  rays  are  gradually  interpolated,  rather 
irregularly,  between  the  older  ones,  all  around  the  circumference,  but 
not  without  some  order.1 

The  same  is  true  of  Labidiaster,  which  has  numerous  long  rays 
when  adult. 

In  Pycnopodia,  the  number  of  rays  when  adult  is  twenty  to  twenty- 
four.  It  normally  starts,  when  very  small,  with  six  equal  rays.  A 
new  pair  of  rays  then  appears,  one  on  each  side  of  one  of  the  primary 
rays ;  then  another  pair  appears  just  back  of  these,  and  so  on  in  suc- 
cessive pairs.2  This  would  always  produce  an  even  number  of  rays. 
But  variations  from  this  bilateral  regularity  often  appear,  producing 
odd  numbers.  (See  pis.  LXXIII  and  LXXIV.) 

1  See  H.  L.  Clark,  Starfishes  of  the  Genus  Heliaster,  Bull.  Mus.  Comp.  Zool., 
vol.  LI,  pp.  25-76,  pis.  i-viii,  1907.    In  this  paper  all  the  species  are  fully  de- 
scribed and  their  modes  of  adding  new  rays  are  explained. 

2  See  plates.    See  Professor  W.  E.  Ritter  and  G.  R.  Crocker  (Proc.  Wash- 
ington Acad.  Sci.,  vol.  n,  pp.  247-274,  1900),  who  give  a  good  account  of  the 
process. 


SHALLOW- WATER   STARFISHES  13 

All  the  known  species  of  these  three  remarkable  genera  are  from 
the  Pacific  coast  of  America  and  the  outlying  islands.1 

Their  resemblance  to  certain  fossil  starfishes  of  the  Devonian  is 
very  striking,  and  may  indicate  a  continuous  descent  from  those 
ancient  forms.  If  so,  it  would  be  good  proof  of  the  antiquity  of  the 
West  Coast  forms,  as  mentioned  above. 

Aside  from  these  remarkable  and  localized  genera,  there  are  other 
genera  of  the  fauna  that  have  numerous  rays.  The  genus  Solaster 
is  represented  there  by  five  or  six  shallow-water  species,  and 
Crossaster  by  one.  All  these  are  multiple-rayed,  the  rays  varying 
from  eight  to  fifteen.  In  these  genera,  there  is  no  evidence  of  the 
interpolation  of  new  rays  in  post-larval  life.  They  are  also  genera 
that  are  widely  distributed  in  all  seas.  (See  pis.  viu  and  x.) 

They  appear  to  be  more  numerous  on  the  Northwest  Coast  than 
elsewhere,  both  as  to  individuals  and  species.  Several  additional 
species  occur  in  deeper  water  off  the  coast.  The  group  may  have 
originated  in  the  North  Pacific,  in  former  geologic  ages. 

The  genus  Pteraster  in  all  other  seas  has  five  rays,  except 
P.  obscurus,  of  the  North  Atlantic  and  Arctic,  which  ordinarily  has 
six  or  seven.  Variety  octaster  V.,  of  Bering  Sea,  normally  has  eight 
rays,  but  sometimes  seven.  Mr.  Fisher  thinks  that  it  is  only  a  variety 
of  P.  obscurus,  but  even  if  so,  it  is  singular  that  the  latter  should  be 
six-rayed  in  the  Atlantic  and  anywhere  from  six-rayed  to  eight- 
rayed  in  the  North  Pacific. 

So,  likewise,  other  genera  that  are  generally  five-rayed  in  all  other 
parts  of  the  world  are  apt  to  have  six-rayed  species  on  the  North 
Pacific  coast.  This  is  true  of  the  genus  Asterias  and  its  subdivisions, 
in  which  we  find  many  six-rayed  species  on  that  coast,  as  well  as 
numerous  five-rayed  ones.  Some  of  those  that  are  ordinarily  five- 
rayed  are  also  often  found  with  six  or  seven  rays.  This  occurs 
in  other  regions,  but  not  so  frequently  as  there. 

Among  the  species  of  Asteriidae  having  normally  six  or  more 
rays,  the  following  may  be  mentioned :  Pisaster  grandis  (St.),  a  very 
large  species;  Orthasterias  merrianti  V. ;  A.  acervata  St.;  A.  kath- 
erince  Gray ;  A.  multiclava  V. ;  Stenasterias  macropora  V. ;  Leptas- 
terias  hexactis  (St.)  ;  L.  vancouveri  (Per.)  ;  L.  aqualis  (St.)  ;  L. 
coei  V. ;  L.  epichlora  (Br.),  subsp.  alaskensis  V.,  and  several  varie- 
ties of  the  same  species;  and  L.  macouni  V. 

Some  of  these  have  occurred  with  seven  rays,  and  some  with  five, 
as  abnormal  variations. 

1  Two  species  of  Heliaster  occur  at  the  Galapagos  Islands,  and  one  at  Juan 
Fernandez.    These  were  evidently  derived  from  Pacific  coast  ancestors. 


14  VERRILL 

The  following  species,  which  are  normally  five-rayed,  have  been 
found  with  six  rays: 

Henricia  leviuscula  (ST.),  from  Alaska. 

Henricia  sanguinolenta  (Minx.),  Bering  Sea. 

Pisaster  ochraceus  (En.),  Monterey,  Calif.,  and  British  Columbia. 

Patiria  miniata  (BR.).    Six-rayed  common;  four- rayed  and  seven-rayed  rare; 

California. 
Dermasterias  imbricata  (GR.),  Sitka. 

When  we  consider  the  great  geological  antiquity  and  remarkable 
persistence  of  the  five-rayed  condition  in  echinoderms  generally, 
it  is  very  remarkable  that  so  many  genera  and  species  of  existing 
starfishes  should  have  acquired  the  peculiarity  of  having  higher 
numbers.  It  seems  natural  to  conclude  that  there  must  be  some 
special  advantage  gained  by  this  increase  in  rays,  which  does  not 
obtain  in  the  other  classes  to  the  same  extent. 

At  the  present  time  all  known  species  of  sea-urchins  and  holo- 
thurians  are  normally  five-rayed,  and  the  same  holds  good  through 
all  the  geologic  ages  for  the  former  class  at  least.  Abnormal  sea- 
urchins  with  four  and  with  six  rays  have  been  observed  in  a  number 
of  species.  Dr.  Robert  T.  Jackson,  in  his  recent  extensive  work,1 
has  discussed  this  and  various  other  variations  of  sea-urchins  very 
fully.  He  states  that  he  has  personally  studied  seventy-one  cases 
of  more  or  less  complete  variations  from  the  perfect  five-rayed  condi- 
tion. To  find  these  required  the  special  examination  of  50,000  speci- 
mens, including  many  species. 

In  some  species  about  one  individual  in  a  thousand  shows  varia- 
tions of  this  kind.1 

The  serpent-stars  (Ophiuroidea)  show  more  variation  in  the 
number  of  rays  than  the  sea-urchins  do.  Several  genera  contain 
some  species  that  are  normally  six-rayed.  The  common  Ophiocoma 
pumila  of  Bermuda  and  the  West  Indies  has  about  equal  numbers  of 
five-rayed  and  six-rayed  individuals  living  together.  Many  of  the 
species  of  Ophiactis  have  six  to  eight  rays  while  young,  and  divide 
autotomously,  but  when  mature  they  are  usually  either  regularly 
five-rayed  or  six-rayed.  Yet  some  of  the  species  of  the  same  genus 
are  always  five-rayed  and  do  not  divide ;  so  it  does  not  seem  to  be  a 
matter  of  great  importance  in  this  genus. 

1  Phylogeny  of  the  Echini,  Mem.  Boston  Soc.  Nat.  Hist,  vol.  x,  1912. 
Variations  from  the  pentamerous  system  are  described  on  pp.  35-50. 

1  See  also  A.  E.  Verrill,  Amer.  Naturalist,  vol.  XLIII,  p.  545,  for  a  discussion 
of  the  subject  with  figures. 


SHALLOW- WATER   STARFISHES  1 5 

Variations  from  the  five-rayed  condition  occur  not  very  rarely 
among  living  Crinoids,  and  were  not  uncommon  among  the  ancient 
fossil  pentremites  and  cystideans;1  but  on  the  whole  the  crinoids 
have  been  pretty  constantly  five-rayed  through  all  the  geologic  ages. 

We  must  conclude  that  all  these  variations  originated  at  first  as 
"  sports,"  which  have  persisted  by  heredity  and  natural  selection, 
because  they  were  advantageous.  It  is  easy  to  conjecture  that,  in 
the  case  of  two  starfishes,  otherwise  similar  in  size  and  structure, 
living  together  on  a  rocky  shore  and  exposed  to  violent  surf,  the  one 
with  six  rays  would  be  able  to  cling  more  securely  to  the  rocks  than 
the  one  with  five  rays.  Therefore,  because  of  the  increased  number 
of  ambulacral  sucker-feet,  it  might  well  be  the  form  preserved  by 
natural  selection,  unless  for  some  other  important  but  unknown 
reason,  the  five-rayed  condition  has  certain  other  more  important 
advantages. 

It  is  certainly  true  that  most  of  the  shallow-water  species  with 
multiple  rays  live  among  rocks  in  situations  exposed  to  the  surf. 
This  is  true  of  the  seven  species  of  Heliaster,  with  very  numerous 
rays;  and  of  Pycnopodia  with  twenty  to  twenty-four  rays,  and  of 
the  various  shallow-water  and  littoral  species  of  Solaster  and  Cross- 
aster,  which  usually  have  nine  to  fifteen  rays  (rarely  eight  or  less). 
It  is  also  true  of  the  numerous  six-rayed  species  of  Asterias, 
Pisaster  and  allied  genera.3 

However,  the  ability  to  cling  tenaciously  to  rocks  may  be  perfected 
in  other  ways,  involving  equally  an  increased  number  of  sucker-feet. 
This  is  often  attained  by  lengthening  the  rays,  as  in  many  species  of 
Asterias;  by  crowding  the  suckers  into  more  than  four  rows,  as  in 
some  large  species  of  Pisaster;  and  by  increasing  the  size  and 
strength  of  the  suckers. 

Although  it  seems  probable  that  the  added  protection  gained 
against  the  violence  of  the  waves  by  the  evolution  of  more  numerous 
rays  and  suckers  is  a  real  cause  for  the  retention  of  this  feature,  it 
may  not  be  the  principal  one.  The  attainment  of  four  rows  of  sucker- 

1  Some  of  the  paleozoic  Cystidea  were  tri-radial.  Hence  it  has  been  thought 
by  some  writers  that  this  was  the  more  primitive  condition  and  that  the 
pentamerous  condition  was  acquired  by  the  addition  of  another  pair  of  rays. 
See  Bather,  op.  cit,  1901. 

*  The  family  Brisingidae,  however,  is  mostly  confined  to  deep  water.  All  the 
species  are  multirayed,  with  long  rays.  Some  of  them  climb  over  gorgonians, 
as  do  many  ophiuroids.  The  genus  Labidiaster,  of  the  Patagonian  region,  is 
found  in  shallow  water  among  rocks  and  sea-weeds.  It  has  numerous  long 
rays. 


1 6  VERRILL 

feet  in  Asteriidae,  as  compared  with  the  more  primitive  two-rowed 
condition,  was  probably  due  to  the  same  causes. 

I  am  inclined  to  believe  that  the  increase  of  rays  has  been  due  more 
to  the  advantage  gained  in  holding  their  food  securely,  and  in 
opening  bivalves,  than  for  holding  to  the  rocks,  though  both  go 
together.  The  starfishes  are  the  most  predaceous  of  the  echinoderms. 
They  feed  largely  on  bivalve  shells  that  cling  firmly  to  the  rocks, 
like  the  oysters,  and  on  others  that  must  be  pulled  open  by  means  of 
their  numerous  muscular  suckers,  part  of  which  must  be  used  for 
keeping  in  a  suitable  position  and  part  for  the  opening  of  the  oyster, 
at  the  same  time,  so  that  the  more  arms  and  suckers  they  have,  the 
more  rapidly  and  effectively  they  can  secure  their  food,  in  case  they 
are  feeding  on  large  bivalves. 

We  must  admit  that,  so  far  as  known,  the  five-rayed  and  six-rayed 
individuals  of  a  species  appear  to  be  equally  well  nourished  and  grow 
to  equal  size.  Also  that  the  normally  six-rayed  species  of  Asterias 
are  commonly  no  larger,  nor  more  robust,  than  the  allied  five-rayed 
species,  in  the  same  environment.  Even  the  four-rayed  individuals 
appear  to  be  well  nourished  and  of  ordinary  size.  It  is  quite  possible, 
however,  that  they  grow  more  slowly.  It  is  known  that  the  rate  of 
growth  of  starfishes  is  very  rapid  when  food  is  abundant,  but  very 
slow  when  food  is  scarce.  No  observations  have  been  made  on  the 
comparative  rate  of  growth  of  six-rayed  specimens. 

It  must  be  remembered  that  the  variation  in  the  number  of  rays 
is  necessarily  attended  by  extensive  changes  in  the  number,  size,  and 
form  of  the  skeletal  plates  of  the  body;  also  in  the  number  of 
ambulacral  feet  and  water  tubes,  nerve  ganglions,  nerve  cords, 
stomach  lobes,  hepatic  glands,  and  all  other  internal  organs.1  A  six- 
rayed  specimen  has  twelve  reproductive  glands,  instead  of  the  ten 
in  its  five-rayed  competitor.  If  the  number  of  ovules  be  propor- 
tionately large,  it  would  produce  twenty  per  cent  more  young. 

So,  likewise,  it  would  have  an  additional  stomach  lobe  and  two 
more  hepatic  glands.  This  would  perhaps  be  of  advantage  in  the 

1  This  does  not  hold  good  in  the  cases  of  the  multirayed  species  of  Heliaster, 
according  to  Clark.  (The  Genus  Heliaster,  1907.)  This  genus  has  but  five 
stomach  lobes  and  five  pairs  of  gastric  retractor  muscles,  whatever  may  be  the 
number  of  rays.  Yet  the  added  rays  contribute  to  digestion  in  that  group, 
for  each  ray  has  a  pair  of  hepatic  glands  and  a  pair  of  gonads. 

The  term  "  hepatic  gland  "  must  not  be  understood  as  implying  that  these 
glands  correspond  precisely  with  the  liver  of  vertebrates,  for  it  is  known  that 
the  digestive  fluids  of  echinoderms  are  different  chemically  from  those  of  the 
latter.  Some  claim  that  these  glands  of  starfishes  are  more  like  the  pancreas 
in  function. 


SHALLOW- WATER   STARFISHES  I/ 

digestion  of  food  and  cause  more  rapid  growth  and  earlier  maturity, 
if  not  greater  reproductive  powers. 

It  would  be  a  very  interesting  experiment  to  try  breeding  six-rayed 
or  seven-rayed  varieties  of  some  common  five-rayed  species. 

I  have  found  five-rayed  young  (about  5  per  cent)  among  those 
carried  by  a  six-rayed  mother,  in  the  case  of  Leptasterias  epichlora. 
Fisher  states  that  he  has  found  seven-rayed  young  in  the  gonocodium 
of  an  eight-rayed  Pteraster  obscurus,  and  one  with  nine  rays.  That 
six-rayed  and  seven-rayed  varieties  might  be  easily  obtained  by 
selection  and  isolation  is  very  probable.  I  have  observed  that  in  the 
case  of  our  common  New  England  starfish  (Asterias  forbesi)  six- 
rayed  and  seven-rayed  individuals  are  much  more  common  in  some 
localities  than  elsewhere,  indicating,  perhaps,  a  tendency  to  the 
inheritance  of  these  features. 

SPECIES,  SUBSPECIES,  AND  VARIETIES. 

Within  the  limits  of  recognized  species,  certain  groups  of  indi- 
viduals, apparently  having  a  definite  distribution  and  more  or  less 
permanency  of  characters,  are  regarded  as  bathymetrical  or  geo- 
graphic races.  Such  races  are  here  called  subspecies. 

In  certain  genera,  for  example  Asterias,  Henricia,  Solaster,  speci- 
mens occur  which  cannot  be  referred  definitely  to  any  species  or  sub- 
species. Some  of  these  appear  to  be  local  variations,  due  to  unfavor- 
able environments ;  "  sports,"  "  freaks,"  or  "  hybrids ; "  others  may  be 
abnormal  individuals.  They  are  here  called  varieties.  If  in  future 
the  characters  of  any  of  these  varieties  prove  fairly  constant,  it  may 
become  necessary  to  elevate  them  to  subspecific  or  even  specific  rank. 
To  determine  the  status  of  these  forms,  much  larger  series  of  speci- 
mens than  are  at  present  available  will  be  necessary. 

It  is  not  unreasonable  to  think  that  some  of  these  "  varieties  "  and 
"  subspecies  "  are  incipient  species,  now  in  the  process  of  evolution, 
and  that  eventually  some  may  become  fixed  species,  while  others  will 
disappear  by  the  elimination  of  the  most  unfit. 

The  Northwest  Coast  appears  to  be  one  vast  nursery  for  the  origi- 
nation and  evolution  of  new  varieties,  subspecies  and  species  of  star- 
fishes and  various  other  groups  of  marine  animals. 

A  peculiar  difficulty  in  the  determination  of  genera  and  species  of 
starfishes  consists  in  the  fact  that  many,  especially  of  the  Asteriidae, 
do  not  attain  their  adult  characters  until  of  considerable  size,  so  that 
young,  or  even  half-grown  specimens,  may  appear  to  belong  to  a 
different  species,  or  even  a  different  genus,  for  the  more  complex 


l8  VERRILL 

species  pass  through  stages  corresponding  to  the  adult  condition  of 
simpler  or  more  primitive  genera,  and  there  is  often  no  way  to  tell 
whether  a  small  specimen  is  mature  or  not,  without  comparison  with 
a  series. 

HYBRIDISM  OF  ASSOCIATED  SPECIES. 

Owing  to  the  fact  that  numerous  related  species  of  several  genera 
are  living  together  on  the  Northwest  Coast,  it  is  natural  to  conclude 
that  some  of  them  may  frequently  hybridize,  especially  if  their 
breeding  seasons  are  coincident.  I  have  seen  many  hybrids  between 
Asterias  vulgaris  and  A.  forbesi  of  the  New  England  coast. 

There  is  considerable  evidence  that  this  often  occurs  on  the  Pacific 
Coast,  leading  to  the  existence  of  many  very  puzzling  specimens  in 
some  groups. 

I  have  personally  seen  a  number  of  apparent  hybrids  between 
diverse  species  of  Asterias,  Pisaster,  Henricia,  and  other  genera 
abundant  on  that  coast.  Perhaps  some  of  the  equally  puzzling  speci- 
mens of  Solaster  may  also  be  hybrids. 

To  determine  this  matter  satisfactorily  requires  large  collections, 
and  especially  observations  made  on  the  living  specimens  in  their 
natural  environment.  The  littoral  species  of  Leptasterias  that  carry 
their  young,  abundant  on  the  shores,  are  eminently  adapted  for  such 
studies,  and  are  easily  raised. 

This  can  best  be  done  at  one  of  the  seaside  biological  stations, 
and  for  that  reason  I  do  not  propose  to  discuss  the  matter  at  length 
in  this  place.  Dr.  Fisher,  also,  has  found  numerous  varieties  and 
connecting  forms,  some  of  which  he  thinks  may  be  hybrids,  among 
the  large  numbers  of  specimens  of  Henricia  that  he  has  studied. 

It  is  quite  probable  that  some  of  the  now  well  established  species 
of  starfishes  of  that  coast  originated  at  first  as  hybrids  and  that 
others  are  now  in  the  process  of  differentiation  into  varieties  and 
species. 

FAUNAL  RELATIONS  AND  DISTRIBUTION  OF  THE  SPECIES. 

Details  of  the  distribution  of  the  species  must  be  left  to  the  end  of 
this  report,  but  there  are  some  interesting  facts  that  may  well  be 
mentioned  here. 

The  shallow-water  starfish  fauna  of  the  Northwest  Coast,  taken 
collectively,  may  be  considered  as  consisting  of  three  parts. 

i  st.  Those  of  more  southern  or  tropical  origin  that  have  migrated 
northward  to  California  or  beyond. 


SHALLOW-WATER   STARFISHES  19 

2d.  Those  that  are  of  Arctic  or  circumpolar  distribution  and 
extend  southward  into  Bering  Sea,  and  in  many  cases  to  the  Aleutian 
Islands,  or  much  farther  south,  especially  in  the  deeper  waters. 
Their  origin  may  have  been  in  the  North  Pacific  originally,  in  many 
cases. 

3d.  Those  that  appear  to  have  originated  in  the  region  extending 
from  the  Aleutian  Islands  to  California. 

The  last  group  is  by  far  the  larger  and  more  important.  Very 
likely  it  is  possible  to  divide  that  very  extensive  coast  line  into  two  or 
more  faunal  districts,  but  for  the  starfishes  this  does  not  seem 
to  be  necessary  for  the  present,  for  many  species  range  throughout 
that  whole  extent  of  coast. 

The  limits  of  distribution  on  that  coast  seem  to  be  determined 
entirely  by  the  temperature  of  the  water,  especially  in  the  breeding 
season,  due  probably  to  the  greater  sensitiveness  of  the  free-swim- 
ming larval  forms.  The  adults  can  regulate  their  temperatures  by 
migrating  into  deeper  or  shallower  water  as  occasions  require. 

Among  the  more  southern  forms  are  Orthasterias  forreri,  Pisaster 
capitatus,  P.  paucispinus,  Marthasterias  sertulifera,  species  of  Astro- 
pecten  and  Luidia,  Linckia  Columbia,  etc. 

Among  the  arctic  and  in  part  circumpolar  species  are  Asterias 
acervata,  A.  multiclava,  L.  arctica,  C.  cribraria,  Allasterias  rathbuni, 
Henricia  sanguinolenta,  H.  tumida,  H.  arctica,  Solaster  endeca, 
Crossaster  papposus,  Pteraster  obscurus,  and  var.  octaster,  P.  mili- 
taris,  Diplopteraster  multipes,  Tosiaster  arcticus,  C.  granularis, 
Leptychaster  arcticus,  Ctenodiscus  crispatus,  etc. 

The  species  apparently  indigenous  on  that  coast  are  too  numerous 
to  enumerate  here.  Some  of  them  are  nearly  allied  to  those  of  the 
North  Atlantic  and  other  regions,  but  many  are  very  peculiar  and 
have  no  near  allies  in  other  regions,  so  far  as  known. 

A  number  of  the  genera  and  higher  groups  are  peculiar  to  that 
coast,  others  have  there  a  remarkable  development  in  number  and 
variety  of  species,  showing  that  their  evolution  must  have  gone 
along  continuously  for  vast  periods  of  time.  In  many  cases  primi- 
tive and  highly  specialized  species  of  a  family  are  found  associated. 
Among  these  peculiar  types  are  Pycnopodia,  Rathbunaster,  several 
species  of  Pisaster,  many  species  of  Asteriinae,  Dermasterias  imbri- 
cata,  Leptychaster  (Glyphaster)  anomalus,  Pteraster  tesselatus, 
Bunodaster  ritteri,  etc. 

Fuller  lists  will  be  given  at  the  end  of  this  work,  with  more 
details  of  their  distribution. 


Class  ASTERIOIDEA. 

MORPHOLOGICAL  FEATURES. 

The  Asterioidea1  have  a  polygonal  or  star-shaped  body,  in  which 
the  rays  are  direct  prolongations  of  the  body  itself,  and  contain  exten- 
sions of  the  body  cavity  and  more  or  less  of  the  viscera,  especially  one 
or  more  pairs  of  gonads  and  a  pair  of  digestive  glands ;  generally, 
also,  a  lobe  of  the  saccular  stomach. 

The  skeleton  is  made  up  of  large  numbers  of  ossicles  or  plates, 
mostly  articulated  so  as  to  be  more  or  less  movable,  giving  flexibility 
both  to  the  rays  and  to  the  disk,  though  in  some  species  (certain 
Goniasteridae)  the  flexibility  is  slight,  except  at  the  tips  of  the  rays. 

The  actinal  side  of  the  disk  and  rays  has  deep  radial  ambulacral 
grooves,  extending  to  the  tip  of  the  rays.  The  roof  of  the  groove 
is  supported  by  the  two  rows  of  ambulacral  ossicles,  arranged  like 
rafters,  or  in  close,  inverted  V-shaped  pairs  of  compressed  plates  or 
bars,  between  which  there  are  rows  of  pores  for  the  passage  of  the 
ambulacral  feet.2 

In  the  middle  line  of  each  ray  and  external  to  the  ambulacral 
plates  are  situated  the  radial  nerve  and  blood-vessel.  The  radial 
water  tube  supplies  water  to  the  locomotive  tubes  through  the 
medium  of  muscular  ampullae,  usually  double,  situated  internally 
above  the  ambulacral  plates.  In  Brisingidae  ampullae  are  lacking; 
in  Echinasteridae  they  are  single. 

There  is  no  median  row  of  calcareous  plates  covering  the  ambu- 
lacral areas  and  radial  nerve  and  blood-vessel,  such  as  exists  in 
Ophiuroidea. 

The  tip  of  each  ray  ends  in  a  special  terminal  ocular,  or  apical, 
plate,  supporting  a  pigmented  ocellus,  to  which  the  radial  nerve 
extends.  According  to  the  studies  of  Fewkes,  1888,  these  are  the 
first  plates  to  appear  in  the  young. 

The  grooves  are  bordered  on  each  side  by  a  row  of  plates  called 
adambulacral,  which  always  bear  spines. 

1  This  spelling  of  the  name  is  preferred  because  it  is  derived  from  Asterias, 
not  from  Aster. 

"In  many  paleozoic  fossil  starfishes  the  ambulacral  plates  are  not  opposite, 
in  pairs,  but  alternate.    This  rarely  occurs  in  existing  species,  though  I  have 
noticed  it  in  Pycnopodia  as  an  abnormal  variation  in  some  of  the  rays,  and 
also  that  it  may  occur  from  lateral  bending. 
(20) 


SHALLOW- WATER   STARFISHES  21 

The  mouth  is  central,  dilatable,  and  surrounded  by  soft  membrane. 
The  so-called  jaws  are  merely  the  adoral  ambulacral  and  adambu- 
lacral  plates,  more  or  less  modified  and  coalesced ;  the  "  teeth  "  are 
only  slightly  modified  adambulacral  spines,  in  this  work  called  peroral 
spines.  (See  text-fig.  3.) 

The  stomach  is  very  saccular  and  usually  evertible.  It  usually  has 
a  lobe  and  a  pair  of  digestive  glands  extending  into  the  cavity  of  each 
ray;  but  in  the  case  of  some  multiple-rayed  species  (Heliaster), 
it  has  lobes  corresponding  only  to  the  primary  five  rays.  In  this 
case  the  five  stomach-lobes  do  not  enter  the  rays,  but  the  pairs  of 
digestive  glands  do.  In  some  slender-rayed  genera,  also,  the  stomach 
is  confined  to  the  disk.  The  intestine  is  usually  nearly  or  quite 
abortive  and  not  functionally  active.  The  so-called  "  anus  "  is  a 
dorsal  pore,  chiefly  for  the  discharge  of  secretions  from  the  dorsal 
glands,  or  "  csecal  appendages,"  probably  nephridial  in  function, 
and  called  nephridial  glands  in  this  work. 

Commonly  there  is  a  single  pair  of  branched  gonads  in  the  proxi- 
mal part  of  each  ray,  with  simple  ducts  discharging  through  a  pair 
of  interradial  pores,  which  may  be  either  ventral  or  dorsal.  In  cer- 
tain Brisingidae,  Luidiidae,  and  in  a  few  other  families  there  are 
several  pairs  of  gonads  and  genital  pores  arranged  serially  along  the 
sides  of  each  ray. 

The  madreporite  or  madreporic  plate  is  dorsal,1  excentric,  and 
commonly  single,  yet  in  some  multiradiate  species  there  may  be  sev- 
eral. It  is  an  organ  primarily  for  the  purpose  of  eliminating  the 
excess  of  absorbed  water  from  the  ambulacral  tubes  and  body-cavity. 

The  sides  of  the  rays  and  disk  are  generally  supported  by  two 
rows  of  marginal  plates,  usually  larger  or  thicker  than  the  other 
plates,  and  commonly  bearing  special  spines.  They  are  called  supra- 
marginals  or  superomarginals  and  inframarginals  or  inferomarginals. 
(See  text-fig.  I.) 

The  upper  row  is  sometimes  much  reduced,  or  obsolete,  and  rarely 
both  rows  are  lacking  or  rudimentary.  These  rows  of  plates  belong 
to  the  primary  system  of  plates,  and  extend  to  the  apical  plate  of 
the  rays,  like  the  ambulacral  and  adambulacral  rows. 

Plates  are  constantly  added  to  these  rows  by  the  budding  in  of 
new  plates  between  the  apical  plate  and  the  one  next  to  it,  the  apical 
plate  being  pushed  farther  outward  and  the  ray  lengthened  at  the 

1  In  some  paleozoic  fossil  starfishes  it  is  said  to  be  ventral,  but  it  is  not  so 
in  any  living  species.  The  statement  by  Gregory  (op.  cit.,  p.  238,  1900)  that  it 
is  ventral  in  Asterina  is  erroneous. 


22  VERRILL 

same  time.  The  first  median  dorsal  plate  of  the  rays  appears  very 
early  in  the  young,  and  the  first  marginals  and  adambulacrals  soon 
after. 

In  many  genera  and  families,  but  not  in  all,  additional  rows  of 
plates  may  be  interpolated  between  the  marginal  rows  proximally, 
and  are  called  intermarginals ;  or  between  the  inferomarginals  and 
the  adambulacrals,  when  they  are  called  interactinals,  intermediate 
actinals,  or  simply  actinal  plates.1  These  do  not  appear  very  early  in 
the  young,  and  are  often  without  spines. 

These  interactinal  and  intermarginal  plates  do  not  belong  to  the 
primary  system  of  plates,  nor  do  they  commonly  reach  the  apical 
plate.  Their  new  plates  develop  mostly  at  the  tips  of  the  rows,  as 
the  starfish  grows  larger,  and  new  rows  may  be  interpolated  till  they 
sometimes  become  very  numerous,  in  order  to  increase  the  size  of 
the  rays. 

The  dorsal  skeleton  of  the  disk,  in  a  five-rayed  species,  primarily 
has  five  basal  radials  (or  first  dorsals),  five  genitals,  and  five  pairs 
of  interradial  plates,  besides  the  plates  that  later  become  the  apical 
or  ocular  plates,  and  the  centro-dorsal  plate.  By  the  interpolation  of 
new  ossicles  and  plates  in  various  ways,  the  structure  often  becomes 
very  complex. 

On  the  rays  we  can  usually  distinguish  a  median  or  carinal  row, 
extending  from  the  basal  radial  to  the  apical  plate.  Other  regular 
rows  may  develop  each  side  of  this  (the  dorso-laterals),  or  the  whole 
surface  may  become  covered  with  a  tesselated  arrangement,  or  a 
reticulated  system  of  plates  and  transverse  ossicles.2 

The  dorsal  plates,  like  the  marginals  and  interactinals,  commonly 
bear  spines  or  small  spinules,  but  they  may  be  covered  with  granules, 
or  with  a  smooth  soft  integument,  or  even  appear  quite  naked,  being 
then  covered  only  with  a  thin  membrane. 

These  plates  and  their  armatures  of  spinules  take  several  special 
names,  according  to  their  forms  and  structure,  and  are  often  char- 
acteristic of  special  genera  and  families  and  higher  groups. 

When  they  become  columnar  or  of  hour-glass  shape,  and  have  the 
summit  covered  with  a  radiating  cluster  of  small  slender  spinules, 

1  For  more  details  of  the  rows  of  plates  and  their  sequence,  see  below, 
under  the  family  Asteriidae,  and  text-figure,  i,  i-iv.  For  the  sequence  in  the 
early  stages  of  the  development  of  the  primary  plates,  see  J.  W.  Fewkes : 
On  the  Development  of  the  Calcareous  Plates  of  Asterias,  1888. 

*For  more  details  of  the  skeletal  plates,  see  the  discussions  under  the  sev- 
eral orders  and  families  below,  especially  under  Forcipulata  and  Phaner- 


SHALLOW-WATER   STARFISHES  23 

they  are  called  paxillse.  These  are  most  typically  formed  in  Astro- 
pecten,  Luidia,  and  allied  genera,  and  are  characteristic  of  the  sub- 
order Paxillosa.1 

Pedicellariae  of  one  or  more  kinds  are  generally  present,  but  are 
entirely  lacking  or  very  rare  in  certain  families,  such  as  the 
Solasteridae,  Echinasteridae,  Pterasteridae,1  and  in  some  genera  of 
other  families.  Their  uses  are  imperfectly  known.  One  use  is  to 
keep  the  body  free  from  dirt  and  parasites." 

Another  use,  perhaps  the  most  important,  seems  to  be,  according 
to  my  own  observations,  to  retain  the  discharged  unfertilized  ova 
until  fertilization  takes  place,  thus  preventing  their  dispersal  and 
loss.  This,  however,  needs  further  investigations  on  living  starfishes. 

These  curious  organs  are  not  found  in  other  classes  of  Echino- 
derms,  except  in  the  Echinoidea.  In  the  latter  they  are  commonly 
mounted  on  long  stalks  and  generally  have  three  valves,  sometimes 
two  to  four  or  even  more. 

In  the  Asterioidea  they  are  usually  sessile  or  (Order  Forcipulosa) 
have  very  short  pedicels,  and  they  are  most  frequently  bivalved. 

They  are  of  several  different  types,  which  are  commonly  char- 
acteristic of  the  orders  and  families,  and  often  of  the  genera. 

Sometimes  there  may  be  bivalved,  trivalved,  four-valved,  and 
five-valved  ones  on  a  single  specimen.  (See  below,  under  Der- 
masterias  imbricata,  and  pi.  vi,  figs.  4,  5  P,  P'.).  In  some  cases  they 
are  not  really  valvular,  but  consist  of  several  movable  spinules, 
arranged  in  convergent  groups  or  opposed  rows  (pi.  xxxiv,  P.  P.). 
The  sessile  valvular  forms  are  generally  situated  over  a  pore,  pene- 
trating a  plate,  for  the  passage  of  nerves,  etc.  (Pis.  XLVII,  XLVIII.) 

The  different  kinds  take  special  names.  For  further  details  of 
the  pedicellariae  and  their  special  names,  see  below,  under  the  sev- 
eral orders,  and  plates  LXXV-LXXXV. 

In  nearly  all  species  there  are  numerous  small,  soft,  tubular  out- 
growths from  the  body-wall,  called  papulae,  serving  for  respiratory 
purposes.  They  may  be  in  large  groups,  especially  between  the 

1  For  various  other  special  forms  and  their  names,  see  below,  under  Phaner- 
ozona  and  Valvulosa. 

2  In  the  last  two  of  these  three  families,  and  perhaps  in  the  first,  the  mother 
carries  and  protects  the  eggs  and  young  until  they  are  able  to  provide  for 
themselves. 

3  The  large  bivalve  pedicellariae  of  Hippasteria  are  very  muscular.    I  have 
lifted  a  large  living  starfish  of  this  kind,  weighing  about  a  pound,  entirely  out 
of  water  by  a  toothpick,  seized  by  the  jaws  of  a  dorsal  pedicellaria. 


24  VERRILL 

dorsal  and  lateral  ossicles,  or  stand  singly,  or  they  may  occur  only  in 
special  areas.  Rarely  they  are  branched. 

Most  shallow-water  starfishes  that  have  been  studied  in  this  respect 
have  a  free-swimming,  bilateral  larval  form,  known  as  a  brachiolaria ; 
yet  a  considerable  number  carry  their  eggs  and  young  till  they 
develop  the  starfish  form,  as  explained  above.  These  have  an  abbre- 
viated metamorphosis.  The  two  methods  may  occur  in  different 
genera  of  the  same  family  (e.  g.,  Asteriidce). 

The  embryology  of  the  deep-sea  species  is  unknown  in  most  cases, 
and  may  be  diverse,  or  even  more  abbreviated. 

CLASSIFICATION  OF  ASTERIOIDEA. 

In  this  report  I  have  adopted  the  division  of  the  class  into  three 
large  orders,  viz. : 

I.  FORCIPULOSA  or  FORCIPULATA. 
II.  SPINULOSA. 
III.  PHANEROZONA. 

The  last  has  two  suborders,  considered  orders  by  some  recent 
writers,  viz. : 

I.  VALVULOSA  or  VALVATA. 
II.  PAXILLOSA. 

Order  FORCIPULOSA  Verrill,  or  FORCIPULATA  Perrier. 

Stelleridce  forcipulatce  PERRIER,  Mem.  Etoiles  de  Mer,  pp.  166,  188,  1876. 

Forcipulata  PERRIER,  Exped.  Sci.  Trav.  et  Talisman,  p.  27,  1894. 

Asteries  Ambulacraires  VIGUIER,  Squellette  des  Stellerides,  Arch.  Zool.  Exper., 

vii,  p.  93,  1878. 

Cryptozonia  (pars)  SLADEN,  Voy.  Chall.,  xxx,  pp.  xxxiv,  397,  1889. 
Adetopneusia  (pars)  and  Leptostroteria  (pars)  SLADEN,  op.  cit,  p.  xxxiv. 

The  form  is  always  stellate,  often  with  long  rays,  commonly  five, 
but  often  multiple.  Ambulacral  plates,  except  the  orals,  are  usually 
short  and  closely  crowded  (leptostroteriaT) ,  but  not  in  Brisingidae. 
The  proximal  pair  is  elongated  and,  except  in  Pedicellaster  and  a 
few  others,  forms  the  inner  end  of  the  jaw.  Ambulacral  feet  are 
generally  arranged  in  four  rows, — but  in  two  rows  in  Pedicellas- 
teridae,  Brisingidae,  and  Zoroasteridae. 

In  some  large  species  of  Pisaster  they  may  form  six  or  more  rows 
by  crowding,  especially  subproximally.  They  are  always  terminated 
by  suckers. 


SHALLOW-WATER   STARFISHES  2$ 

Adambulacral  plates  are  generally  short  and  crowded,  equal  in 
number  to  the  ambulacrals.  They  may  each  bear  a  single  spine 
(monacanthid) ,  or  two  spines  (diplacanthid) ,  or  alternately  one  and 
two  (subdiplacanthid) ,  or  rarely  three  spines  (triplacanthid) . 

The  superior  and  inferior  marginal  plates  are  rather  small 
(cryptozonial) ,  but  are  generally  distinct  and  bear  spines,  often 
longer  than  those  on  the  dorsals.  In  some  Brisingida  one  row,  or 
rarely  both  rows,  may  be  lacking.  They  extend  to  the  apical  plate 
and  increase  in  number  during  the  whole  period  of  growth  by  the 
addition  of  new  plates  next  to  the  apical. 

The  dorsal  skeleton  is  variously  constituted,  but  usually  consists 
of  plates  and  ossicles,  often  very  unequal  in  size  and  form,  so  united 
as  to  move  freely,  leaving  many  papular  areas  between  them.  In 
some  of  the  Brisingidae  and  in  Pycnopodia  the  ossicles  are  mostly 
nearly  abortive,  roundish,  and  isolated  in  the  integument. 

The  plates  generally  bear  spines,  either  singly  or  in  clusters; 
sometimes  they  are  reduced  nearly  to  the  form  of  granules ;  never 
paxilliform.  The  papulae  may  be  few,  or  many  in  large  clusters. 
They  usually  occur  both  on  the  dorsal  and  on  the  ventral  surfaces, — 
adetopneusic  arrangement. 

The  Forcipulosa,  as  the  name  indicates,  are  especially  character- 
ized by  the  presence  of  very  peculiar,  pincer-like,  two-bladed  pedicel- 
lariae,  usually  of  two  forms,  movably  attached  to  the  dermis  of  the 
spines,  or  to  the  surface  between,  by  means  of  longer  or  shorter 
flexible  dermal  pedicels. 

VARIOUS  KINDS  OF  PEDICELLARIJE. 

Except  possibly  in  a  few  rare  cases,  and  in  quite  immature  speci- 
mens, at  least  one  form  of  these  is  always  present. 

Of  the  two  principal  forms,  those  known  as  forcipate  or  "  minor 
pedicellarice"  are  the  most  abundant  and  occur  most  constantly. 
Such  pedicellariae  do  not  occur  on  starfishes  of  any  other  order. 
Those  nearest  analogous  occur  on  certain  Echinoidea. 

In  the  larger  kind,  called  "major  pedicellaria"  by  Stimpson;1 

*I  prefer  to  use  ordinarily  the  names  first  given  by  Stimpson,  in  1861,  for 
the  two  principal  forms  of  pedicellarias,  viz.,  major  pedicellariee  and  minor 
pedicellaria.  These  terms  are  sufficiently  descriptive,  and  not  so  liable  to  be 
confused  as  those  proposed  later  by  Herapath,  though  the  latter  have  some 
advantages.  It  is  desirable,  however,  to  have  some  additional  special  de- 
scriptive terms  for  varieties  of  each  sort.  Either  form  may  be  attached  to 
the  spines  or  to  the  general  integument.  The  minor  pedicellariae  that  sur- 
round spines  in  wreaths,  may  be  called  circumspinal;  those  in  smaller  irregu- 
lar clusters  on  the  spines,  epispinal;  those  attached  to  the  general  surface  of 


26  VERRILL 

"  pe dicellaires  droites"  by  Perrier,  1869;  and  "  forficiform  pedicel- 
larice"  by  Herapath,  1866,  and  by  Sladen,  1889,  the  jaws  are  nearly 
straight,  articulated  at  the  base  by  a  simple  joint  to  a  thick  basal 
piece,  so  that  they  open  and  close  like  forceps.  These  may  be  either 
dermal  or  attached  to  the  spines,  either  singly  or  in  clusters;  they 
often  have  short  pedicels.  In  the  Brisingida  and  in  Pedicellaster 
they  are  usually  lacking.  (See  pi.  XLIX,  figs.  3-30?;  pi.  xxx,  etc.) 

The  second  kind,  called  "  minor  pedicellarice  "  by  Stimpson,  "  pe  di- 
cellaires croises"  by  Perrier,  "crossed  pedicellaria"  by  several 
writers,  and  "  forcipiform  pedicellaria "  by  Herapath,  are  usually 
much  smaller  and  the  blades  are  curved  and  crossed  something  like 
those  of  scissors,  and  so  articulated  that  they  open  and  close  like 
scissors,  tweezers,  or  pincers.  These,  like  the  others,  may  be  attached 
directly  to  the  integument,  either  singly  or  in  clusters,  or  to  the 
spines,  to  the  pedicels  of  the  larger  forms,  to  saccular  dermal 
growths  around  the  spines,  to  the  inner  edge  of  the  ambulacral 
grooves,  or  rarely  even  to  the  sucker-feet.  (See  plates  LXXVI- 
LXXXIV.)  They  are  most  commonly  attached  by  slender  and  some- 
times long  pedicels,  and  aggregated  into  wreaths  or  clusters  on  the 
spines.  They  are  often  so  abundant,  in  the  wreaths  around  the  dorsal 
spines,  that  they  nearly  or  quite  conceal  the  spines  and  integument 
in  living  specimens.  In  some  cases  they  are  attached  in  large  num- 
bers to  dermal  sheaths  or  sacks  loosely  surrounding  the  spines.  They 
often  have  a  formidable  array  of  minute  sharp  denticles,  and  are 
frequently  very  characteristic  of  species.  (See  text-fig.  2.) 

FAMILIES  AND  SUBFAMILIES  OF  FORCIPULOSA. 
This  order  now  includes  the  following  families  and  subfamilies : 
Family  ASTERIIDJE. 
Subfamilies  ASTERIINM;  STICHASTERINJE;  PYCNO- 

PODIIN;E;  HELIASTERIN^E. 

Family  ZOROASTERID2E. 
Family  PEDICELLASTERID&. 
Family  BRISINGIDJE. 

Subfamilies  BRISINGINJE;  LABIDIASTERINJE.   (Type, 
Labidiaster  Liitk.) 

the  body  are  dermal;  those  on  a  papular  area  are  papular;  those  on  the 
adambulacral  spines  or  plates  are  adambulacral.  Major  pedicellariae  may  also 
frequently  occur  within  the  adambulacral  grooves,  attached  to  the  inner  sur- 
face of  the  adambulacral  plates,  where  they  may  be  called  intra-adambulacral. 


SHALLOW-WATER   STARFISHES  27 

Family  ASTERIID2E  Gray  (emended). 

Asteriidce  GRAY,  Ann.  Mag.  Nat.  Hist.,  vi,  p.  178,  1840;  Synopsis,  p.  I,  1866. 
Perrier,  Revis.  Stell.,  Arch.  Zool.  Exper.  et  Gen.,  iv,  p.  302,  1875;  Mem. 
Etoiles  de  Mer,  pp.  167,  198,  1876.  Viguier,  Squelette  des  Stellerides,  pp. 
93,  99,  pl-  v,  figs,  i-io,  11-12,  1878. 

Asteriida  (emended)  -f-  Stichasterida  SLADEN,  Voyage  Chall.,  xxx,  pp.  430, 
560,  1889.  Perrier,  Exp.  Trav.  et  Talisman,  pp.  43,  105,  107,  128,  1894; 
Contrib.  1'etude  des  Stellerides  Atlan.  Nord,  pp.  25,  31,  1896. 

v/ 
Rays  five  to  twenty  or  more,  but  most  frequently  five  or  six. 

Madreporic  plate  generally  single  in  five-rayed  and  six-rayed  forms, 
but  often  two  or  more  in  those  with  a  variable  number  of  rays. 
Dorsal  and  actinal  plates  various  in  form  and  arrangement.  Odonto- 
phore  usually  formed  of  a  single  piece.  Oral  ambulacra!  plates 
elongated;  those  along  the  grooves  become  very  short  and  com- 
pressed. Papulae  occur  both  on  the  actinal  and  dorsal  sides.  Ambu- 
lacral  feet  usually  crowded  in  four  rows,  sometimes  more.  Pedicel- 
lariae  of  two  kinds  are  present.  Dorsal  ossicles  generally  bear 
spines  or  spinules,  various  in  size  and  kind,  and  sometimes  granules. 
They  are  never  true  paxillae. 

The  mouth  is  large;  the  stomach  saccular  and  evertible;  intestine 
rudimentary  or  abortive.  A  subcentral,  dorsal  pore  (nephridial  pore) 
is  present,  through  which  is  discharged  the  secretion  of  the  branched 
or  lobular  "  rectal "  or  nephridial  glands,  which  are  usually  unequal 
and  often  but  two  or  three  in  number.  This  pore  is  usually  called  the 
"  anal  pore,"  but  rarely,  if  ever,  functions  as  an  anus  in  this  family. 
The  intestine  is  nearly  or  quite  abortive,  in  most  cases. 

I.  MORPHOLOGY  OF  THE  OSSICLES. 

Aside  from  the  ambulacral  and  adambulacral  plates,  the  skeleton 
of  the  rays  consists  of  five  fundamental  or  primary  rows  of  ossicles, 
viz.,  the  median  dorsal  or  carinals;  the  two  superomarginals;  and  the 
two  infer omarginals.  The  latter  may  or  may  not  be  confined  to  the 
ventral  side.  (See  fig.  i,  i.) 

The  ossicles  of  adjacent  rows  may  be  articulated  directly  by  their 
lobes,  or  either  large  or  small,  simple  connecting  ossicles  may  inter- 
vene. 

This  simple  or  primitive  type  of  skeleton  is  found  in  Heterasterias 
volsellata  (Sla.),  and  in  a  few  other  species,  especially  when  young. 

In  most  cases  more  or  less  numerous  ossicles  or  rows  of  ossicles 
are  interpolated  during  growth  between  the  five  primary  rows,  either 
above  or  below,  or  between  the  marginals,  thus  giving  rise  to  many 


VERRILL 


md 


. 


FIG.  i. 


Diagrams  generalized  to  illustrate  progressive  ontogenetic  and  phylog_enetic  development 
of  the  plates  and  spines  in  the  rays  of  starfishes  (Asteriidee).  Lettering  as  follows;  the 
primary  rows  are  shaded: 

md,  Median  dorsal  or  carinal  row.  m.  Marginal  rows,  m',  Intermarginal  row.  _  sm, 
Supermarginal  row.  im,  Inferomarginal  row.  ad,  Adambulacral  row.  din,  Dorsal-inter- 
mediate, medio-lateral,  or  dorso-lateral  rows,  p  or  pa,  Peractinal  row  of  plates.  sa,  Sub- 
actinal  rows.  These  and  the  peractinals  are  the  interacting  plates,  collectively,  o,  The 
ocular,  apical,  or  terminal  plate. 

i,  The  simplest  or  most  primitive  condition,  common  in  the  young;  uncommon  in  the 
adults,  in  which  only  the  primary  rows  of  plates  are  developed. 

The  longitudinal  area  between  ad  and  the  next  row  is  called  the  perambulacral  lane  or 
channel;  the  space  between  the  marginals  (m)  is  called  the  intermarginal  lane  or  channel; 
the  area  next  above  the  upper  marginal  is  called  the  supramarginal  lane  or  channel.  These 
"  lanes  "  or  "  channels  "  are  always  crossed  by  transverse  connective  ossicles  or  by  over- 
lapping lobes  of  the  plates,  dividing  them  up  into  larger  or  smaller  papular  areas,  and  they 
usually  bear  more  or  fewer  of  the  larger  pedicellarise  in  the  Asteriidx. 

ii,  The  condition,  more  advanced,  in  which  the  peractinal  row  (p)  and  one  medio-lateral 
row  (din)  are  developed.  A  common  condition  in  the  Asteriinse.  This  is  called  the  mono- 
actinoplacid  condition. 

iii,  A  more  advanced  condition  in  which  an  additional  interactinal  row  (sa,  subactinal) 
and  a  second  dorso-lateral  or  medio-lateral  row  Win)  have  appeared. 

iv,  A  more  complex  state,  in  which  there  are  two  subactinal  rows  (ja),  and  an  inter- 
marginal row  («'). 

This  is  a  common  condition,  but  more  medio-lateral  rows  are  often  added,  as  well  as  more 
subactinals,  in  such  genera  as  Pisaster,  Evasterias,  etc.  This  is  called  the  polyactinoplacid 
condition.  All  these  plates  usually  bear  spines,  one  or  more  each.  They  may  be  easily 
visible  or  they  may  be  entirely  concealed  by  a  thick  tough  skin.  The  peractinal  or  sub- 
actinal plates,  when  thin  or  small,  may  be  destitute  of  spines  and  stand  edgewise,  so  as  to 
show  only  the  edge  at  the  surface,  or  they  may  be  so  closely  joined  to  the  inferomarginals 
as  to  require  cleaning  with  potash  to  be  seen. 


SHALLOW-WATER   STARFISHES  29 

different  styles  of  skeleton  in  the  more  complex  genera.     (Fig.  i, 
ii-iv.) 

The  carinal  or  median  dorsal  plates  (wrf)  usually  remain  distinct, 
but  are  often  small  and  not  easily  recognizable.  The  superomar- 
ginals  (sin)  are  nearly  always  easily  recognizable,  though  small,  and 
they  generally  bear  spines  larger  than  the  dorsals.  They  never  form 
a  stout  margin,  as  in  the  Paxillosa. 

The  inferomarginal  plates  (im,  fig.  i)  are  generally  very  evident 
and  carry  one  or  more  rows  of  spines  usually  longer  or  larger  than 
the  dorsals.  They  are  usually  separated  from  the  superodorsals  by  a 
more  or  less  wide,  naked  lateral  lane  or  channel  on  which  are  rows 
of  papular  groups,  and  frequently  many  of  the  larger  pedicellariae. 
In  some  cases  one  or  more  short  intermediate  rows  of  lateral  plates 
(intermarginals,  w'),  bearing  spines,  may  be  interpolated  in  these 
areas,  proximally. 

Both  marginal  rows  normally  reach  the  ocular  or  apical  plate.  The 
peractinal  row  (fig.  i,  pa)  rarely  actually  touches  the  apical 
plate,  and  generally  falls  considerably  short  of  it. 

This  affords  a  useful  character  for  distinguishing  the  inferomar- 
ginal from  the  peractinal  plates.1  This  name  is  proposed  for  the  first 
or  primary  row  of  actinal  plates. 

To  designate  collectively  the  rows  of  plates  that  may  exist  between 
the  inferomarginals  and  the  adambulacrals,  I  propose  to  use  the  term 
" ' interactinals"  in  place  of  the  longer  one,  "intermediate  actinals," 
used  by  Sladen  and  others.  I  would  do  away  with  the  use  of  the 
words  "  actinal  spines  "  or  plates  used  by  many  writers  as  including 
the  inferomarginals  with  the  true  actinals.  Ventrals  may  be  less 
objectionable  if  such  a  collective  name  be  needed,  and  is  so  used  by 
Perrier  and  others. 

The  upper  marginal  row  of  ossicles  is  usually  pretty  clearly 
defined  by  special  spines.  This  superomarginal  row  usually  diverges 
from  the  lower  one,  or  turns  upward  at  the  base  of  the  rays,  and 
may  run  a  little  upon  the  dorsal  surface  to  join  its  counterpart  on 
the  next  ray  at  the  dorsal  interradial  angle  or  axil.  The  plates  here 
called  subactinals  (text-fig,  i,  so)  appear  later  than  the  peractinals. 
They  are  often  lacking  in  the  young,  up  to  30  to  50  mm.  in  diameter, 
even  when  they  are  present  in  the  adult.  They  often  extend  only  on 
part  of  the  ray,  proximally.  The  number  of  rows  of  these  in  large 
species  increases  with  the  age. 

1  It  seems  desirable  to  have  special  and  convenient  terms  to  designate  the 
several  series  of  plates  and  spines  in  this  and  related  families. 


3O  VERRILL 

The  row  of  actinal  plates  that  unites  with  the  adambulacrals,  when 
there  are  several  rows,  may  be  called  synactinal,  to  indicate  its  func- 
tion. It  may  consist  either  of  the  peractinals  or  of  subactinals. 

When  these  rows  are  very  much  alike  in  size  and  form,  mistakes 
have  often  been  made  in  descriptions,  the  peractinal  row  having 
often  been  described  as  the  inferomarginal  row  or  vice  versa,  or  else 
no  distinction  has  been  made  by  the  earlier  writers. 

The  most  positive  criterion,  in  doubtful  cases,  is  to  trace  these 
rows  of  plates  to  their  origin.  True  marginal  rows  originate  at  the 
ocular  plate;  peractinals,  when  present,  often  terminate  near  the 
ocular ;  but  other  rows  generally  fail  to  reach  the  end  of  the  ray. 

The  inferomarginal  plates  may  be  directly  joined  to  the  adambu- 
lacrals by  a  lobe  or  apophysis,  but  more  commonly  there  is  at  least 
one  row  of  actinal  plates  between,  and  sometimes,  besides  this  first 
or  peractinal  row  (pet),  there  may  be  one  or  more  shorter  rows  of 
subactinal  plates  (sa)  interpolated  proximally.  All  these  plates 
usually  bear  spines,  but  in  some  cases  all  the  interactinal  plates  are 
rudimentary  and  spineless. 

The  secondary  dorsal  skeleton  varies  much,  but  is  most  commonly 
chiefly  made  up  of  unequal  and  more  or  less  imbricated  ossicles  or 
lobulated  plates,  with  the  ends  or  lobes  of  adjacent  plates  over- 
lapping, so  as  to  leave  rather  wide  papular  spaces  between  them. 
Or  the  plates  may  be  broader,  somewhat  rhombic,  lobed,  and  more 
closely  imbricated,  as  in  Stichaster.  In  most  cases,  however,  they 
are  also  united  transversely  by  smaller  connective  ossicles. 

II.  PEDICELLARI^E  OF  THE  ASTERIID^E. 
Text-figure  2. 

Pedicellariae  of  both  kinds  are  probably  always  present  in  the 
normal  adults,  though  on  some  individuals  they  may  be  few  or 
lacking. 

The  major  or  forficulate  pedicellariae  are  generally  ovate  or  lanceo- 
late, sometimes  wedge-shaped,  stone-hammer-shaped,  lyrate,  or 
spatulate.  The  tips  may  be  acute,  plain,  flat,  or  denticulate. 

They  are  mostly  dermal,  but  may  also  occur  on  the  ventral  and  oral 
spines,  or  along  the  inner  edges  of  the  grooves,  or  attached  to  pedi- 
cels in  clusters  within  the  grooves,  or  even  on  the  pedicels  of  the 
ambulacral  feet.  They  may  be  of  several  sizes  and  forms  on  one 
specimen. 

Certain  species  of  Stichaster,  Coronaster,  etc.,  have  a  peculiar  very 
large  form  of  dermal  pedicellarise,  often  as  thick  as  the  spines,  in 


SHALLOW-  WATER   STARFISHES  31 

which  the  broad  valves  terminate  in  a  series  of  curved  denticles,  so 
that  when  closed  the  valves  resemble  a  pair  of  clasped  hands,  or 
better  still,  a  pair  of  clasped  feet  of  a  cat,  with  the  claws  protruded. 
These  may  be  called  felipedal  or  unguiculate.  In  many  other  cases 
similar  gigantic  pedicellariae  have  spatulate  valves,  with  a  smooth 
or  denticulate  edge.  In  Pisaster  they  are  sessile,  stout,  erect,  stone- 
hammer-shaped  or  wedge-shaped,  with  serrate  or  unguiculate  edges. 
(See  pi.  XLIX,  figs.  3-36.) 

These  peculiar  giant  forms  are  usually  accompanied  by  others  of 
the  ordinary  ovate  form,  much  smaller  in  size,  and  they  are  usually 
characteristic  of  special  generic  groups.  But  as  such  groups  are 
sometimes  widely  separated,  both  morphologically  and  geographi- 
cally, it  is  not  unlikely  that  these  are  survivals  of  a  type  of  pedicel- 
lariae that,  at  some  remote  period,  was  common  to  the  whole  family. 

The  minor  or  forcipulate  pedicellariae  (text-fig.  2)  generally  form 
wreaths  or  large  or  small  clusters  on  the  spines,  but  they  may 
also  occur,  either  singly  or  in  clusters,  on  the  integument  of  the 
dorsal  or  lateral  plates,  on  the  papular  areas,  and  in  other  situations. 

III.  RELATIVE  IMPORTANCE  OF  MORPHOLOGICAL  CHARACTERS 

IN 


Perrier,  Sladen,  Bell,  and  others  who  have  divided  this  family, 
have  differed  considerably  in  their  estimation  of  the  relative  impor- 
tance of  the  various  morphological  characters. 

A.    MODES  OF  GROWTH  AND  INCREASE  IN  NUMBER  OF  SKELETAL  PLATES. 

Text-figure  I,  i-iv. 

The  number  of  ossicles  and  plates  of  nearly  all  kinds,  except  the 
primary  radials  and  orals,  and  the  oculars,  increase  continually  dur- 
ing the  entire  growth  of  most  starfishes.  The  size  and  age  have  no 
definite  limit,  so  that  unusually  large  specimens  of  any  species  may 
often  occur  with  correspondingly  increased  numbers  of  plates  and 
spines.  Therefore  the  exact  number  of  plates,  say  of  marginals,  is 
of  no  great  importance,  except  when  specimens  of  identical  sizes  or 
ages  are  compared.  In  the  early  stages  of  growth,  the  first  dorsal 
plates  that  appear  are  the  central,  primary  interradials,  the  radials, 
and  the  ocular  plates.  A  little  later  the  median  dorsal  radials  and  the 
marginals  appear;  and  the  commencement  of  a  row  between  the 
upper  marginals  and  median  row  often  appears  at  the  same  time  or  a 
little  later.  The  number  of  the  plates  in  all  the  primary  rows  is 


32  VERRILL 

increased  by  the  formation  of  new  plates  between  the  apical  terminal 
or  ocular  plate  and  the  plate  next  to  it,  the  oculars  being  forced 
farther  away  as  the  numbers  and  growth  of  the  plates  continue  to 
increase.  But  besides  the  primary  rows  of  plates,  other  rows  and 
scattered  ossicles  may  appear,  to  fill  up  the  dermal  spaces  produced 
by  the  increasing  diameter  of  the  rays  and  disk.  In  some  forms  of 
starfishes  this  is  accomplished  largely  by  the  rapid  growth  of  the 
primary  plates,  which  may  thus  become  very  large,  but  this  produces 
a  rather  inflexible  skeleton.  In  the  Asteriidse  flexibility  seems  to 
be  an  essential  feature,  especially  as  most  of  the  species  cling  to 
irregular  stones  and  similar  objects,  and  rarely  live  habitually  on 
smooth  mud  or  sand.  Therefore,  the  skeleton  has  come  to  be  com- 
pleted in  this  group  by  the  constant  addition  of  small  ossicles  movably 
articulated  between  the  rows  of  larger  ones,  especially  on  the  dorsal 
surface,  both  longitudinally  and  transversely. 

These  later  interpolated  ossicles  may  form  regular  or  irregular 
longitudinal  rows  between  the  median  and  marginal  rows,  or  they 
may  be  so  interpolated  as  to  form  a  more  or  less  regular  reticu- 
lation ;  in  other  cases  they  may  be  entirely  irregular,  with  no  deter- 
minate arrangement,  seeming  to  be  introduced  wherever  needed  at 
the  time. 

These  variations  in  the  modes  of  interpolation  of  the  plates, 
together  with  their  forms,  which  may  vary  from  linear  to  broad, 
angular,  lobate,  and  scale-like  forms,  combined  with  variations  in 
the  spines,  give  rise  to  the  great  variety  of  structures  and  forms 
seen  in  the  dorsal  surface  of  starfishes  of  this  family. 

Moreover,  in  most  species,  the  rapid  increase  of  the  dorsal 
skeleton  alone  apparently  does  not  give  sufficient  space  for  the 
rapidly  growing  internal  organs,  especially  the  reproductive  organs, 
within  the  bases  of  the  rays,  and  therefore  new  rows  of  plates  (fig. 
i,  iii,  iv)  must  be  interpolated  between  the  inferomarginals  and  the 
adambulacrals,  and  sometimes  between  the  upper  and  lower  mar- 
ginals, to  increase  the  diameter  of  the  rays.  The  most  constant  of 
these  rows  is  that  which  I  have  designated  as  the  peractinal  (pa}. 
This  row  appears  very  early  in  many  species  and  often  nearly 
reaches  the  tip  of  the  ray,  but  usually  not  quite  to  the  ocular  plate. 
But  the  other  actinal  rows  (subactinals  or  interactinals)  (sa)  are 
successively  shorter  and  often  do  not  reach  the  middle  of  the  ray. 
The  plates  situated  proximally  in  these  rows  appear  earliest,  and 
new  ones  are  added  at  the  distal  end  of  each  row.  This  is  also  the 
case  with  the  interpolated  dorsal  and  lateral  rows.  In  many  species, 


SHALLOW-WATER   STARFISHES  33 

as  in  those  of  Pisaster,  the  number  of  interpolated  interactinal  rows 
(so)  goes  on  increasing  during  the  whole  or  most  of  the  period  of 
growth,  so  that  in  large  specimens  there  may  be  four  or  five  rows  on 
each  side.  But  in  many  other  species  the  number  of  rows  is  fixed 
very  early,  the  young,  two  inches  in  diameter,  having  as  many  rows 
as  those  six  to  eight  inches  in  diameter,  the  subsequent  increase  in 
size  being  due  to  the  enlargement  of  the  plates  and  the  formation 
of  new  ones  dorsally  and  distally.  But  in  many  species  with  long, 
slender  rays  the  peractinal  plates  alone  are  developed,  and  in  a  few 
even  these  are  lacking,  or  else  so  small  as  not  to  be  visible  exter- 
nally, as  in  Urasterias  linckii,  etc.  So,  likewise,  in  other  long-rayed 
species,  the  dorsal  skeleton  may  lack  more  than  one  pair  of  interpo- 
lated rows  of  ossicles,  as  in  Coscinasterias. 

B.      INTERACTINAL  PLATES  AND  SPINES. 

Text-figure  I. 

The  presence  or  absence  of  actinal  plates  and  spines,  and  whether 
one  row  or  several  rows  of  these  plates  be  developed,  are  characters 
that  often  seem  to  be  of  generic  value.  Those  forms  that  have  no 
such  plates  at  maturity,  or  have  only  one  imperfect  row,  are  probably 
the  more  primitive  types,  or  at  least  they  have  probably  inherited  and 
preserved  this  primitive  character.  The  very  young  starfish  in  all 
the  genera  apparently  has  none  of  these  plates,  but  acquires  them 
as  growth  proceeds.  They  are  lacking  or  rudimentary  in  Urasterias, 
Distolasterias,  Stylasterias,  and  some  other  groups.  They  exist  in 
only  one  row  in  Orthasterias,  Coscinasterias,  etc.  In  true  Asterias 
and  some  other  groups,  one  or  two,  or  more,  incomplete  subactinal 
rows  are  usually  added.  In  Evasterias,  Pisaster,  and  Cosmasterias, 
two,  three,  or  more  rows  are  present.  In  those  species  that  have 
subactinal  plates,  these  increase  in  number  during  the  whole  period 
of  growth,  and  new  rows  may  also  be  added  continually.  Never- 
theless these  plates  seem  to  afford  morphological  characters  of  much 
value  in  classification,  and  they  are  usually  coincident  with  other 
special  features.  It  is  of  great  importance  that  correct  distinctions 
should  be  made  between  the  actinal,  inferomarginal,  and  supramar- 
ginal  plates  and  spines.  This  has  not  been  done  in  most  of  the 
earlier  descriptions.  Side  views  of  the  rays  are  necessary  to  show 
these  plates  properly,  but  such  views  are  rarely  given.  In  many 
species  they  are  without  spines  and  so  concealed  by  thick  skin  that 
they  cannot  be  determined  without  preparation,  and  therefore  are 
not  visible  in  alcoholic  specimens. 
4 


34  VERRILL 

C.    FORM   AND  ARRANGEMENT  OF  THE  DORSAL  OSSICLES. 

The  form  and  arrangement  of  the  dorsal  ossicles  vary  considerably 
and  often  afford  characters  of  much  value.  Aside  from  the  stichas- 
terial  arrangement,  there  are  two  main  types, — the  reticulate  and 
the  longitudinal, — in  both  of  which  the  principal  plates  are  lobed, 
substellate,  or  cruciform.  The  former,  which  is  seen  in  true 
Asterias,  Pisaster,  etc.,  includes  several  subtypes, — as  the  stellate- 
reticulate,  seen  in  Pisaster  fissipinus;  the  areolate,  seen  in  E.  tro- 
schelii;  and  the  irregular  or  indeterminate,  seen  in  Asterias  vulgaris, 
A.  rub  ens,  etc.  These  variations  depend  partly  upon  variations  in 
the  forms  of  the  plates  and  partly  on  their  arrangement  and  the 
existence  of  supplementary  ossicles,  and  especially  on  the  size  and  the 
form  of  the  papular  areas. 

The  abactinal  skeleton  of  the  longitudinal  type  may  consist  of 
only  three  row's  of  stout  dorsal  plates,  besides  the  marginals,  with 
few  or  no  supplementary  ossicles,  as  in  Coscinasterias;  or  the 
median  may  alone  remain  distinct,  as  in  some  species  of  Urasterias. 
But  in  other  related  groups  there  may  be  five  or  more  regular  rows. 
These  last  have  but  three  rows  when  quite  young,  so  that  the  forms 
with  three  rows  are  probably  more  primitive.  Urasterias  may  be 
still  more  primitive.  Its  skeleton  is  very  likely  a  degenerate  type. 
In  the  stichasterial  arrangement  the  plates  become  broader,  more 
angular,  and  less  lobed,  and  they  are  usually  united  directly  together 
by  their  overlapping  or  adjacent  edges  in  rather  regular,  longitu- 
dinal, imbricated  rows ;  or  they  may  have  a  tesselated  arrangement. 
The  intervening  papular  areas  are  consequently  small  and  rather 
regularly  arranged. 

D.  DORSAL  SPINES;  FORM  AND  ARRANGEMENT. 

The  number,  form,  and  arrangement  of  the  dorsal  spines  vary 
widely.  Their  arrangement,  when  coincident  with  that  of  the  plates, 
is  a  matter  of  much  importance,  but  widely  different  appearances 
may  occur  in  a  single  species,  due  to  the  variable  number  of  spines 
that  may  occur  on  a  single  plate,  combined  with  their  variations  in 
size  and  shape.  No  character  is  more  likely  to  lead  to  mistakes  in  the 
determination  of  species  and  genera  than  the  appearance  of  the 
dorsal  spines,  especially  in  those  groups  which,  like  restricted 
Asterias,  have  very  numerous  spines  with  an  indeterminate  or 
irregular  arrangement.  But  in  forms  that  have  a  limited  number  or 
regular  arrangement  of  plates,  the  spines  often  conform  strictly  to 


SHALLOW-WATER   STARFISHES 


35 


the  plates,  each  plate  bearing-  one  or  two  spines,  and  then  they  may 
be  very  reliable  guides,  in  case  the  specimens  are  adult.  The  number 
of  plates  or  spines  in  a  row,  and  often,  also,  the  number  of  rows, 
increase  with  age.  The  supplementary  or  connective  ossicles,  when 
present,  also  increase  with  age,  and  may  bear  corresponding  spines. 
Mr.  Bell,  in  his  arrangement  (1881,  p.  502),  made  a  special  dis- 
tinction between  those  species  that  bear  certain  spines  on  isolated  or 
special  dermal  ossicles  (autacanthid)  and  those  that  bear  spines  only 
on  the  regular  skeletal  ossicles  (typacanthid} .  But  the  special  der- 
mal ossicles,  at  least  in  most  cases,  seem  to  be  merely  new  skeletal 
ossicles  in  process  of  formation,  which  are  destined,  a  little  later, 
to  become  articulated  with  the  older  ossicles.  Hence  this  character 
seems  to  be  of  little  importance  and  variable  according  to  the  age  of 
the  specimens,  in  most  species. 


FIG.  2. 

Minor  pedicellarise  of  Pycnopodia  helianthoides  (a,  fr);  profile  views.     X  146. 

The  dorsal  and  marginal  spines  may  be  similar  or  dissimilar.  The 
spines  may  be  large  and  long,  or  slender,  short  and  thick,  or  capitate. 
They  may  stand  singly  on  the  plates,  or  in  groups,  or  they  may  be 
reduced  to  the  form  of  small  spinules  covering  the  plates  in  large 
numbers,  or  even  nearly  to  granules,  but  they  do  not  form  paxillae 
nor  pseudopaxillae. 

E.    PEDICELLARLE  J  SPECIAL  FORMS. 

Peculiar  or  unusual  forms  of  major  pedicellarise  occur  on  many 
species,  and  sometimes  species  of  the  most  diverse  structures  and 
belonging  to  widely  separate  faunae  will  agree  in  having  some  of 
the  felipedal  type,  mentioned  above.  Such  special  forms  of  pedicel- 
lariae  are  doubtless  of  great  value  for  indicating  community  of  origin 
and  generic  affinity,  when  they  occur  on  species  that  are  otherwise 
structurally  similar ;  but  they  must  not  be  made  to  overshadow  other 
structural  features,  for  that  would  lead  to  absurd  results,  as  bringing 


36  VERRILL 

together  such  unlike  forms  as  Coronaster  of  the  North  Atlantic  and 
Stichaster  of  the  South  Pacific — forms  that  evidently  belong  to  dis- 
tinct families.  In  the  case  of  Pisaster,  our  eight  species,  though  so 
different  in  their  dorsal  spines,  all  have  similar  and  peculiar  erect, 
stout,  unguiculate  pedicellariae  of  a  type  rarely  to  be  found  in  other 
groups. 

Some  of  the  remarkable  forms  of  pedicellariae  are  probably  sur- 
vivals of  such  structures  present  in  remote  ancestors,  common  to  this 
and  the  allied  families  in  former  geologic  periods,  rather  than 
structures  independently  developed  in  diverse  genera,  now  living  in 
different  oceans.  Their  complexity  of  structure  seems  to  be  too 
great  to  have  arisen  independently  in  modern  times. 

F.    MADREPORITE  OR  MADREPORIC  PLATE,  AND  SURROUNDING  SPINES. 

Several  writers  have  made  considerable  use  of  the  character  of 
this  plate.  In  some  of  the  species  having  more  than  six  rays,  or  a 
variable  number,  there  may  be  two  or  more  madreporites.  Such 
species  are  also,  in  most  cases,  subject  to  fission,  as  in  Coscinasterias, 
Stephanasterias,  etc.  The  number  of  madreporites  is  variable  in 
each  species  of  this  kind,  while  closely  related  species  may  have  but 
one.  The  madreporite  is  sometimes  surrounded  by  a  definite  circle 
of  special  spines  (echinoplacid  Bell),1  but  in  closely  related  species 
no  such  regular  circle  of  surrounding  spines  exists. 

In  some  cases  different  specimens  of  the  same  species  may  vary 
in  this  respect.  Therefore  it  cannot  be  considered  a  character  of 
more  than  specific  value,  and  sometimes  not  even  varietal.  Bell,  in 
his  arrangement,  appears  to  have  given  too  much  importance  to  it. 
The  same  remarks,  as  to  variability,  apply  to  a  naked  groove  or 
narrow  channel  that  sometimes  surrounds  the  plate.  The  relative 
position  of  the  plate  and  the  number  and  arrangement  of  its  gyri 
vary  in  different  species,  and  also  greatly  with  age,  but  are  of  some 
specific  value  in  certain  cases,  as  is  also  its  color,  in  life. 

G.    JAWS  AND  ORAL  SPINES. 

Text-figure  3. 

The  jaws  in  this  family  consist  of  an  apical  jaw-plate,  composed 
of  the  first  and  second  pairs  of  adambulacral  plates  closely  united 

1  Among  the  northwestern  American  species  that  are  usually  "  echinoplacid  " 
are  L.  aqualis,  A.  katherince,  A.  acervata,  L.  epichlora  (variable),  L.  coei, 
E.  troschelii  (variable),  etc.  In  general,  species  with  small  and  numerous 
spines  are  apt  to  have  this  character.  It  is  particularly  conspicuous  in  Cosmas- 
terias  lurida. 


SHALLOW-WATER   STARFISHES 


37 


together  and  usually  bearing  six  oral  spines.  The  four  terminal 
spines  (peroral  spines  or  papilla)  are  directed  more  or  less  hori- 
zontally over  the  mouth  (a,  of)  ;  the  two  outer  ones  (a')  are  diver- 
gent and  smaller  and  are  sometimes  lacking.  The  outer  angle  of  the 
jaw,  formed  by  the  second  pair  of  adambulacral  plates,  bears  an  erect 
and  usually  longer  pair  of  (epioral)  spines  (e,  e). 

The  characters  of  these  oral  spines  and  plates  often  afford  good 
specific  distinctions,  and  in  some  cases  at  least  are  of  generic  value ; 
but  they  have  not  yet  been  described  or  figured  with  care,  except  in 
a  comparatively  small  number  of  species,  so  that  at  present  their 


ii 


FIG.  3. 

Orthasterias  calif  arnica  V.,  type,     i,  A  jaw;  a.  a;  a',  a',  apical  or  peroral  spines  with  pedi- 
cellariae  (p,  p);  e,  e,  epiorals;  e' ' ,  odd  epioral;  X  15;  ii,  an  adambulacral  spine.     X  15. 

value  is  somewhat  uncertain.  In  general  characters  they  are  remark- 
ably constant  throughout  this  family,  and  in  species  that  differ  widely 
in  other  respects.  This  would  indicate  that  their  special  arrangement 
has  persisted  for  a  very  long  period  of  time,  for  it  must  have  been 
established  before  the  world-wide  dispersal  of  the  family  took  place, 
and  probably  before  the  family  itself  was  differentiated. 

In  the  calcareous  peristomal  ring  the  jaw-plates,  formed  by  the 
union  of  two  ambulacral  plates,  are  usually  more  prominent  than  the 
plates  alternating  with  them,  known  as  the  odontophores,  formed  by 
the  union  of  two  interambulacral  plates.  Such  an  arrangement  is 


38  VERRILL 

called  an  " adambulacral  jaw"  and  it  is  characteristic  of  the  family 
Asteriidae  and  of  most  other  families  of  Forcipulosa. 

External  to  this  proper  jaw-plate  are  a  series  of  two  to  ten  or 
more  pairs  of  adoral  adambulacral  plates,  each  pair  closely  pressed 
together  (contingent),  to  form  the  adoral  carina,  which  may  be  more 
or  less  compressed  and  thin,  and  is  longest  in  those  species  with  a 
large  disk,  like  the  Pisasters,  in  which  it  often  contains  ten  or  eleven 
pairs  of  united  plates.  The  spines  (adorals)  borne  on  these  plates 
are  often  longer  than  those  beyond. 

H.   ADAMBULACRAL  SPINES  J  ARRANGEMENT. 

Much  importance  has  been  given  by  many  writers  to  the  number  of 
adambulacral  spines  on  each  of  the  plates.  Many  species  have  regu- 
larly a  single  spine  on  each  plate,  thus  forming  a  simple  regular  row 
(monacanthid) .  Others  have  regularly  two  spines  to  a  plate,  form- 
ing two  rows  (diplacanthid) .  But  still  more  frequently  the  spines 
stand  irregularly,  one  or  two  to  a  plate,  or  alternately  one  and  two, 
( subdiplacanthid) .  In  these  the  spines  often  appear  to  form  three 
rows.  In  Allasterias  Ver.  the  inner  spine  is  situated  higher  up  within 
the  groove,  on  alternate  plates. 

In  some  cases,  as  in  Pisaster  and  Coscinasterias,  the  monacanthid 
condition  is  associated  with  other  important  characters,  and  in  these 
it  seems  to  be  really  of  generic  value ;  but  it  should  not  be  used  as  a 
character  of  primary  importance,  nor  by  itself,  for  generic  divisions, 
for  in  some  cases  closely  related  species,  or  individuals  of  the  same 
species,  may  differ  in  this  respect.1  Yet  it  is  a  character  ordinarily 
early  fixed  in  the  young  starfish,  and  therefore  important. 

I.    NUMBER  OF  RAYS;   VARIABILITY. 

The  number  of  rays  can  only  be  used  for  subordinate  purposes, 
for  it  is  apt  to  vary  sporadically,  even  in  species  that  are  the  most 
constant,  like  Asterias  vulgaris  and  A.  forbesi  of  the  Atlantic  coast, 
in  which  we  find,  when  vast  numbers  are  examined,  that  a  certain 
per  cent  will  have  six  rays,  and  a  smaller  per  cent  seven  rays  and 
four  rays,  while  eight-rayed  and  even  nine-rayed  examples  may  occur 
very  rarely;  but  those  cases  with  seven  and  more  rays  may,  in  most 
cases,  be  due  to  abnormal  repairs  after  injury. 

In  certain  species  of  Leptasterias  that  carry  their  eggs  and  young 
till  they  reach  the  period  of  bottom  life,  the  young  carried  by  the 

lfThis  is  true  of  Coscinasterias  tenuispina  and  C.  acutispina  (Stimp.). 


SHALLOW-WATER   STARFISHES  39 

same  parent  may  have  different  numbers  of  rays.  Thus  those  carried 
by  six-rayed  parents  of  L.  epichlora  alaskensis,  were  found  to  be 
mostly  six-rayed ;  but  about  five  per  cent  were  five-rayed,  and  a  few 
were  four-rayed.  (See  below,  and  pi.  LXXXV,  figs.  2,  a-/.) 

In  other  species  and  genera  the  number  of  rays  is  extremely 
variable,  as  in  Coscinasterias,  where  they  vary  indifferently,  from  six 
to  eleven  or  more.  In  Pycnopodia,  additional  rays  bud  in  between 
the  older  ones  in  pairs  during  growth,  so  that  the  adult  may  have 
twenty  to  twenty-four  rays.  This  variation  is  complicated  in  most 
species  of  the  former  group,  and  in  Stephanasterias  albula,  by  their 
remarkable  habit  of  spontaneous  fission  (autotomy)  and  subsequent 
irregular  replacement  of  the  lost  rays  by  each  half.  ( See  pis.  LXXIII 
and  LXXIV.) 

J.    ARRANGEMENT  OF  THE  AMBULACRAL  FEET  OR  SUCKER-TUBES. 

These  are  generally  arranged  in  four  crowded  rows  in  each 
groove,  or  sometimes  in  two  zigzag  rows,  one  on  each  side  of  the 
median  line.  But  in  many  of  the  larger  species  with  broad  rays,  as 
in  the  genus  Pisaster,  they  may  form  six  or  even  eight  rows,  espe- 
cially subproximally.  Pycnopodia  has  four  rows  of  sucker-feet  sub- 
proximally,  and  two  rows  proximally  and  distally  when  adult. 

K.  MODES  OF  DEVELOPMENT;  BROODING  OF  THE  YOUNG;  POSITION  OF 
THE  GENITAL  PORES. 

Two  widely  diverse  methods  of  development  occur  in  the 
Asteriidae,  as  described  above  (p.  7).  So  far  as  now  known,  those 
genera  that  carry  or  brood  their  eggs  and  young  (padophoric  species} 
also  have  their  genital  pores  situated  on  the  actinal  side,  near  the 
mouth ;  while  those  that  disperse  their  minute  eggs,  and  whose  young 
undergo  a  prolonged,  free-swimming  larval  existence,  have  the  geni- 
tal pores  in  the  dorsal  interradial  areas.  It  is  probable  that  this  dif- 
ference is  the  most  important  morphological  character  by  which  the 
family,  or  at  least  the  Asteriinae,  can  be  divided  into  large  generic 
groups.  Unfortunately  this  feature  has  not  been  noted  nor  studied 
except  in  the  case  of  a  small  number  of  the  genera  and  species,  so  that 
at  present,  it  can  be  utilized  only  in  a  limited  number  of  genera. 

CLASSIFICATION  OF  ASTERIID^. 

As  defined  above,  the  Asteriidae  would  include,  as  subfamilies, 
Asteriinae,  Stichasterinae,  Pycnopodiinae,  and  Heliasterinae=Helias- 
teridae  Sla. 


4O  VERRILL 

Subfamily  HELIASTERINJE. 

Subfamily  Heliasterinee  VIGUIER.     Family  Heliasteridce  Perrier;  Sladen. 

The  Heliasterinae  do  not  seem  to  me  to  be  worthy  of  more  than 
subfamily  rank.  They  agree  with  Asteriidae  in  nearly  all  important 
structural  characters.  They  differ  mainly  in  having  very  numerous 
short  rays  and  a  very  broad  disk,  but  in  these  respects  they  are 
closely  approached  by  Pycnopodia  and  Ccelasterias.  Their  dorsal, 
ventral,  and  ambulacral  skeletons  are  distinctly  like  Asteriidae.  The 
only  distinctive  characters  of  much  value  are  the  restriction  of  the 
stomach  to  the  disk  by  a  discobrachial  septum,  and  the  existence  of 
double  septa  between  the  rays. 

The  group,  which  includes  only  the  genus  Heliaster,  is  confined  to 
the  Pacific  coast  of  America  and  the  adjacent  islands,  from  Chile  to 
Lower  California.  (See  H.  L.  Clark,  op.  cit.,  1907,  plates  i-viu,  for 
details  of  structure  and  descriptions  of  all  the  species.) 

Subfamily  PYCNOPODIINJE. 
For  the  characters  of  this  subfamily,  see  below,  p.  197. 

Subfamily  STICHASTERIN1E. 

The  most  developed  forms  of  Stichasterinse,1  e.  g.,  Stichaster 
striatus  —  aurantiacus,  have  several  regular  rows  of  interactinal 
plates ;  other  forms  have  only  one  or  two  rows.  Some  of  the  As- 
teriinae  (Calasterias,  Pisaster,  etc.)  also  have  from  three  to  six  rows 
of  such  plates.  On  the  other  hand,  some  forms  of  Stichasterinae 
(Stichastrella  rose  a,  Stephanasterias  albida,  and  Granaster)  have 
the  dorsal  ossicles  more  or  less  irregularly  arranged ;  and  even  the 
median  row  may  not  be  distinct  in  S.  albula.  In  the  latter  the  ven- 
tral plates  bear  longer  spinules.  Coelasterias  V.  is  an  intermediate 
genus  with  multiple  rays. 

The  Stichasterinae  have,  at  present,  no  known  representatives  on 
the  northwestern  coast,  unless  we  so  reckon  the  Stephanasterias 

1  Stichaster  M.  and  Tr.  (April,  1840)  was  monotypic.  Its  type  was  5".  stria- 
tus M.  and  Tr.  =  Asterias  aurantiaca  Meyen  (not  of  Linne)  =  Tonia  atlantica 
Gray  (Sept.,  1840).  Tonia  is  a  complete  synonym  of  Stichaster,  which  has 
about  six  months  priority.  The  European  species,  A.  rosea,  usually  referred 
to  Stichaster,  and  taken  as  its  type  by  many  writers,  is  a  distinct  genus  which, 
so  far  as  I  know,  has  no  valid  name.  It  may  be  called  STICHASTRELLA, 
5".  rosea  being  the  type.  It  has  two  or  three  rows  of  interactinal  plates; 
adambulacral  plates  bear  two  or  three  spines  irregularly.  Carinal  rows  of 
plates  and  spinules  are  usually  distinct,  but  only  by  the  slightly  larger  size 
of  the  plates. 


SHALLOW-WATER   STARFISHES  4! 

albula?  which  is  a  very  widely  distributed  North  Atlantic  and  Arctic 
species,  which  has  been  found  at  moderate  depths  off  the  northern 
Alaskan  coasts.  I  consider  it  one  of  the  Asteriinae,  allied  to  Lept- 
asterias. 

It  is  autotomous  while  young,  with  a  variable  number  of  rays,  but 
when  large  it  is  usually  regularly  six-rayed. 

The  consideration  of  the  importance  of  the  stichasterial  or  imbri- 
cated arrangement  of  the  dorsal  ossicles  early  led  to  the  separation  of 
Stichaster  as  a  genus,  and  later  as  the  type  of  a  family,  but  perhaps 
even  that  character  does  not  always  indicate  close  affinity  nor  an 
identical  origin,  for  it  may  have  been  developed  independently  in 
different  regions  and  in  distinct  lines  of  descent.  As  a  matter  of 
fact,  we  find  the  greater  number  of  the  species,  and  those  that  are 
the  larger  and  more  typical,  living  in  the  Antarctic  seas  and  on  the 
southwestern  coast  of  South  America,  while  only  a  few  rather  small 
and  less  characteristic  forms  exist  in  northern  seas. 

It  is  not  unlikely  that  the  northern  forms,  like  5.  rosea  and  5. 
albula,  originated  entirely  independently  of  those  of  South  American 
seas,  and  should,  therefore,  be  classed  as  distinct  genera.  This  is 
indicated,  moreover,  by  their  morphological  characters.  *$".  albula, 
especially,  differs  but  little  from  some  forms  of  Lepasterias,  and  may 
well  have  been  developed  from  some  member  of  that  or  a  similar 
group.  Its  embryology  is  not  known. 

However,  all  recent  investigators  admit  that  the  stichasterial 
arrangement  of  the  dorsal  ossicles  is  at  least  of  generic  value/ 

Other  variations  in  these  plates  may  be  of  equal  value. 

The  discovery  of  many  new  generic  and  specific  types  intermediate 
between  typical  Stichaster  and  Asterias,  as  already  intimated,  renders 
it  difficult  to  define  the  limits  of  the  two  so-called  families,  typi- 
fied by  these  genera. 


is  the  Asterias  albula  (Stimp.)  =  Stichaster  albulus  Ver.  =  Stephan- 
asterias  albula  Ver.  =  Stichaster  albulus  S\zden  =  Nanaster  albulus  Perrier, 
1894,  p.  131.  It  is  found  from  the  Arctic  Ocean  to  South  Carolina  and  Georgia, 
in  deep  water,  on  the  east  American  coast,  and  as  a  littoral  species  it  occurs 
as  far  south  as  the  Bay  of  Fundy.  It  is  also  common  on  the  northern  Euro-; 
pean  coasts. 

2  Sladen  has  even  gone  so  far  (op.  cit,  p.  xxxvi,  1889)  as  to  widely  separate 
Stichasteridie  and  Zoroasteridae  from  the  Asteriidae  on  this  character,  almost 
exclusively,  placing  them  between  Linckiidse  and  Solasteridae  —  families  that  do 
not  belong  to  the  Forcipulata.  This  seems  to  me  a  very  unnatural  arrange- 
ment, due  to  overestimating  the  importance  of  the  character  of  the  dorsal 
ossicles. 


42  VERRILL 

Sladen  and  Perrier  disagree  both  as  to  the  character  and  limits  of 
the  two  groups,  because  certain  genera  and  species  present  characters 
of  both  groups,  or  have  intermediate  characters.  Therefore  it 
seems  necessary  to  reunite  them  in  one  family.  However,  it  may 
be  convenient  to  retain  the  groups  as  subfamilies  or  sections  with  no 
very  definite  limits. 

A  pseudostichasterial  condition  is  sometimes  produced  by  the 
existence  of  clusters  of  spinules  in  regular  rows,  on  the  transverse 
dorsal  connective  ossicles.  This  cannot  always  be  determined  with- 
out removing  the  investing  integument. 

Perrier  considered  the  characters  of  the  dorsal  spinules  as  of  para- 
mount importance  in  this  group.  He  would  place  all  the  genera  with 
distinct  spinules  or  spines  in  Asteriidse,  and  those  with  granule-like 
spinules  in  Stichasteridae ; l  but  in  fact  there  are  all  intermediate 
gradations  in  the  character  of  the  armature,  and  it  is  often  difficult 
to  say  whether  these  structures,  in  some  species,  should  be  called 
"  granules  "  or  "  spinules."  This  seems  to  me  a  character  only  of 
specific  value.* 

Subfamily  ASTERIIN&. 

Asterndce  (restr.)   SLADEN,  op.  cit,  430,  560,  1889.     Perrier,  1894,  pp.  105, 
128;  1896,  pp.  25-31. 

As  here  limited  this  subfamily  corresponds  nearly  with  the  re- 
stricted family  Asteriidae  in  the  systems  of  Sladen  and  of  Perrier. 

The  Asteriinae3  are  chiefly  characterized  by  the  more  or  less 
openly  reticulated  arrangement  of  the  dorsal  and  lateral  ossicles, 
especially  of  the  dorsal  ossicles,  which  are  usually  more  stellate  or 
lobulate  and  more  slender  than  in  the  Stichaster  group,  and  generally 
united  either  by  the  ends  of  the  lobes  or  apophyses,  or  else  by  means 
of  interpolated  ossicles,  thus  allowing  more  flexibility  in  the  rays,  and 
larger  papular  areas. 

The  Stichasterinae,  on  the  contrary,  are  chiefly  characterized  by  the 
more  or  less  regular  subtesselated  or  imbricated  arrangement  of  the 

1 S.  gracilis  and  5".  albula,  with  other  related  forms,  have  definite  spines. 

"The  genus  Calvasterias  Perrier,  1895,  has  about  five  series  of  lobed  and 
imbricated,  flat  dorsal  plates,  bearing  few  small  short  spines,  and  entirely 
covered  with  a  thick,  naked,  soft,  canaliculated  dermis. 

The  character  of  the  plating  should  cause  it  to  be  placed  in  the  Sti- 
chasterinae. 

It  is  monacanthid  and  there  is  a  row  of  small  spiniferous,  interactinal  plates, 
at  least  in  C.  stolidota  Sla.,  of  Chile  and  the  Falkland  Islands. 

*  This  form  of  spelling  the  word  seems  necessary,  because  it  is  derived  from 
Asterias,  not  from  Aster. 


SHALLOW-WATER   STARFISHES  43 

ossicles,  both  on  the  sides  and  dorsal  surface  of  the  rays,  the 
ossicles  themselves  being  larger,  broader,  and  more  rhombic,  and 
usually  united  in  more  definite,  longitudinal  rows,  with  smaller  and 
more  regular  intervening  papular  areas.  In  either  group,  as  at 
present  understood,  the  interactinal  or  ventral  plates  may  have  the 
regular  serial  and  subtesselated  arrangement. 

A.  GENERIC  SUBDIVISIONS  OF  ASTERIIN^. 

The  genus  Asterias,  in  the  wider  sense,  as  still  used  by  many 
writers,  includes  a  very  large  number  and  a  great  variety  of  species, 
found  in  all  seas.  Various  attempts  have  been  made  to  divide  the 
genus  into  a  number  of  genera  or  subgenera  on  structural  char- 
acters, but  hitherto  there  has  been  no  general  agreement  among 
writers,  as  to  the  number  or  limits  of  most  of  the  subdivisions.  Nor 
do  I  flatter  myself  that  my  own  views  will  be  altogether  acceptable. 

One  of  the  best  known  classifications  hitherto  proposed  is  that  of 
Sladen.1 

He  admitted,  in  the  restricted  family  Asteriidae  (our  Asteriinae), 
five  distinct  genera,  including  Pycnopodia.  Under  Asterias  he  had 
six  subgenera.  Most  of  these  are  undoubtedly  worthy  of  generic 
rank.  One  of  them  (Leptasterias}  had  been  proposed  as  a  genus  by 
the  present  writer,  many  years  earlier,  and  another  (Stolasterias)  is 
essentially  identical  with  our  Coscinasterias,  1867.  His  subgenus 
Hydrasterias  is  evidently  a  distinct  genus,  peculiar  to  the  deep  seas, 
and  has  already  been  so  recognized  by  Perrier  and  myself.  Perrier, 
in  a  later  work,2  recognized  all  of  Sladen's  divisions  as  genera,  but 
with  the  limits  modified  in  some  cases,  and  proposed  four  more 
generic  subdivisions,  some  of  which  seem  to  have  no  great  syste- 
matic value.  This  is  particularly  true  of  Diplasterias,  separated 
from  Asterias  mainly  on  account  of  the  two  regular  rows  of  adambu- 
lacral  spines, — a  character  that  is  variable  in  this  group  and  often  of 
no  more  than  specific  importance,  taken  alone.  Many  species  of 
Asterias  have  alternately  one  or  two  adambulacral  spines,  and  others 
have  irregularly  one  and  two  to  a  plate.  C.  tenuispina  is  generally 
strictly  monacanthid,  but  unusually  large  specimens  usually  bear  two 
spines  on  a  few  of  the  plates  proximally. 

Perrier,  in  the  article  referred  to,  recognized  fifteen  genera  in  the 
family. 

1  Voyage  Challenger,  Zool.,  xxx,  pp.  560-564,  1889. 

2Exped.  Scientif.  du  Travailleur  et  du  Talisman,  Echinodermes,  pp.  108, 
109,  1894. 


44  VERRILL 

One  of  his  diplacanthid  genera  (Podasterias)  had  P.  liitkeni  Per. 
(not  Asterias  liitkenii  Stimpson)  as  its  type,  and  is,  therefore,  of 
special  interest  in  this  connection.  In  a  subsequent  work *  Perrier  has 
added  two  more  genera:  Distolasterias,  for  D.  stichantha  (Sladen), 
separated  from  Stolasterias  mainly  on  account  of  its  two  rows  of 
adambulacral  spines ;  and  Sclerasterias,  for  S,  guernei  Per.,  a  new 
and  peculiar  type.  But  in  this  later  scheme  he  has  judiciously 
omitted  Diplasterias  ( Perrier,  non  Kcehler,  nee  Ludwig) . 

After  a  careful  study  of  a  large  number  of  species,  belonging  to 
these  various  divisions,  I  am  convinced  that  several  other  genera  or 
subgenera,  based  on  structural  characters  of  quite  as  much  impor- 
tance, can  be  distinguished. 

As  understood  in  this  work,  the  genus  Asterias  (type  A.  rubens 
L.)  will  be  used  in  a  sense  much  more  restricted  than  by  Sladen.  I 
propose  to  keep  separate,  as  genera,  most  of  the  groups  called  sub- 
genera  by  Sladen — as,  for  example,  Cosmasterias,  Smilasterias, 
Coscinasterias,  Hydrasterias — and  most  of  the  groups  more  recently 
proposed  by  Perrier. 

Some  of  these  generic  groups  need  to  be  modified  in  definition  or 
extent,  and  others  should  be  added.  (  See  below,  under  Geographical 
Distribution,  for  the  characters  of  antarctic  genera.) 

I  am  well  aware  that  many  students  of  these  starfishes  do  not 
think  it  worth  while  to  subdivide  Asterias,  even  to  the  extent  that 
was  done  by  Sladen  and  by  Perrier.  But  in  the  case  of  so  extensive 
a  group,  with  a  world-wide  distribution,  it  will  be  practically  impos- 
sible to  intelligently  study  or  discuss  its  evolution  and  distribution 
unless  we  recognize  the  relationship  between  allied  species  by  means 
of  distinctive  group-names  for  minor  groups,  recognizable  by  mor- 
phological characters.  We  should  aim  at  a  phyllogenetic  classifica- 
tion, and  this  is  more  likely  to  be  attained  by  the  recognition  of  mor- 
phologically similar  groups  of  species  than  by  an  indiscriminate 
lumping  of  all  these  diverse  forms  in  one  great  genus,  containing 
perhaps  two  hundred  species  or  more. 

B.  SUBDIVISIONS  OF  THE  ASTERIINJE  HAVING  DEFINITE  LON- 
GITUDINAL ROWS  OF  DORSAL  PLATES  AND  SPINES. 

Among  the  numerous  species  in  which  the  dorsal  plates  and  spines 
have  a  constant  and  characteristic  arrangement,  there  are  several 
divisions,  apparently  of  generic  and  subgeneric  value.  Several  of 

1  Contr.  a  1'etude  des  Stellerides  de  TAtlantique  Nord,  Resultats  des  Cam- 
pagnes  Scientif.,  faits  par  Albert  I,  Prince  de  Monaco,  fas.  xi,  pp.  34,  35,  1896. 


SHALLOW-WATER   STARFISHES  45 

these  are  not  yet  known  from  the  northwestern  coast,  but  are  likely 
to  occur  hereafter,  when  more  dredging  shall  have  been  done,  and 
therefore  it  may  be  useful  to  give  here  a  summary  account  of  the 
principal  groups  of  this  kind. 

Genus  Coscinasterias  Verrill. 

Coscinasterias  VERRILL,  Trans.  Conn.  Acad.  Sci.,  i,  p.  248,  1867.     (Type,  C. 

muricata  V.  =  (?)   C.  calamaria  (Lam.).) 
Stolasterias  (pars)  SLADEN,  Voyage  ChalL,  vol.  xxx,  pp.  563, 583,  1889.    (Type, 

C.  tenuispina.)     Non  Perrier. 
Coscinasterias  and  Polyasterias  PERRIER,  Exped.  Trav.  et  Talisman,  p.  108, 

1894. 

Body  small,  rays  elongated,  somewhat  angular,  usually  carinate. 
Dorsal  ossicles  of  the  rays  stout,  four-lobed,  usually  arranged  in 
three  or  five  regular  rows,  besides  the  upper  marginals,  which  form 
regular  lateral  rows.  Actinal  ossicles  usually  consist  of  one  primary 
(peractinal)  row,  like  the  lower  marginals,  but  they  may  be  rudi- 
mentary and  spineless  in  half-grown  specimens,  and  only  bear  spines 
in  large,  mature  individuals,  as  in  C.  tenuispina. 

Minor  pedicellariae  are  abundant.  Large  forficulate  or  major  pedi- 
cellariae  of  the  usual  forms  are  present,  often  in  considerable  num- 
bers, on  the  dorsal  and  lateral  plates,  and  smaller  ones  occur  on 
the  marginal  areas.  Large  unguiculate  pedicellariae  are  lacking. 
Adambulacral  plates  monacanthid  or  nearly  so.  Often  autotomous; 
rays  variable  in  number.  Madreporites  often  two  or  more. 

Stolasterias,  which  was  proposed  as  a  subgenus  by  Sladen  in  1889, 
was  nearly  identical  with  Coscinasterias,  characterized  by  me  in  1867. 
He  gave  no  reason  for  changing  the  name.  It  should  be  regarded  as 
a  synonym  of  the  latter  and  be  eliminated.  Sladen's  type  was 
A.  tenuispina,  which  is  a  Coscinasterias  with  the  peractinal  plates 
rudimentary  and  without  spines,  except  in  the  larger  specimens. 

M.  Perrier  (1894)  correctly  retained  Coscinasterias  for  the  typical 
forms,  like  calamaria,  but  separated  those  species  that  are  known  to 
undergo  spontaneous  fission  under  the  name  of  Polyasterias  = 
typical  Stolasterias  Sla.,  though  they  do  not  appear  to  differ  much 
in  structure  from  the  preceding,  while  he  retained  Stolasterias  Sla., 
in  a  restricted  sense,  for  those  allied  to  glacialis,  although  C.  tenui- 
spina was  named  by  Sladen  as  the  type. 

As  C.  tenuispina  often  divides  spontaneously,  it  belongs  to  Poly- 
asterias in  Perrier's  arrangement.  This  would  make  the  latter 
strictly  synonymous  with  Sladen's  typical  Stolasterias,  which  should 


46  VERRILL 

be  restricted  to  this  type  as  a  subgenus,  if  retained  at  all.  The 
group  including  glacialis  had  also  been  previously  named  Marth~ 
asterias  by  Jullien  (Bull.  Zool.  Soc.  France). 

No  species  of  typical  Coscinasterias  is  known  to  me  from  the 
northwestern  coast,  but  a  species  (C.  acutispina  Stimp.,  sp.),  similar 
to  C.  tenuispina,  and  often  having  some  diplacanthid  adambulacral 
plates,  occurs  at  Ousima  Island,  Japan.  It  is  autotomous. 

C.  TABLE  OF  EXTRALIMITAL  SPECIES  OF  COSCINASTERIAS 
AND  CLOSELY  ALLIED  FORMS. 

A.  More  than  one  distinct  row  of  dorsal  spines  (usually  three  rows). 

B.  One  row  of  peractinal  spines  and  plates  well  developed.    Rays  variable, 

six  to  twelve.     Mostly  fissiparous. 

C.  Rays   mostly   seven   to   twelve.     Adambulacral    spines    strictly    uniserial 

(monacanthid).    Lower  marginals  bear  two  spines  each.    Usually 
two  or  more  madreporic  plates  are  present. 
C.  muricata   (VER.)    1867.     (J  =  C.  calamaria.)  Tt*^ 
C.  calamaria  (LAM.).    Australia;  Indian  Ocean. 
C.  jehennesii   (PER.).    Madagascar.     (?  =  calamaria,  teste  Sladen.) 
C.  echinata  (GRAY).  Valparaiso.    Rays  eight 
C.  gemmifera  (PER.).     Chile;'  Fiji  Is.     (?  =  echinata.) 
C.  (f)aster  (GRAY).    Rays  twelve  to  thirteen.    Five  rows  of  dorsal 

spines  (Gray). 

CC.  Some  of  the  adambulacral  plates  diplacanthid.    Rays  variable. 
C.  acutispina  ( STIMP.).     Ousima  Island. 
C.  tenuispina  LAM.     (Large  specimens.) 

BB.  No  interactinal  spines ;  peractinal  plates  rudimentary  or  lacking. 
AA.  Only  one  row  of  large  dorsal  plates  and  spines.     Adambulacral  plates 

monacanthid. 

b.  Rays  mostly  five  to  nine.    Fissiparous.    Subgenus  Stolasterias  (Sla.  restr.). 
C.  (S.)  tenuispina  (LAM.)   when  young.     Eastern  Atlantic;  Brazil; 

West  Indies. 

Variety  atlantica  (VER.).    Bermuda;  Cuba, 
bb.  Rays  constantly  five  or  six.    Type  not  fissiparous. 

Marthasterias  (JuL.)  when  young.    Type,  M.  glacialis  (MULLER). 

D.  OBSERVATIONS  ON  VARIOUS  GENERA. 

Genus  Heterasterias  Verrill,  nov. 
Type,  Stolasterias  volsellata  Sla.,  Philippines. 

The  rudimentary  state  of  the  dorsal  skeleton  and  the  lack  of 
peractinal  plates  in  the  named  species  may  entitle  it  to  be  con- 
sidered the  type  of  a  new  generic  division,  as  suggested  by  Sladen. 
This  view  is  strengthened  by  reason  of  the  presence  of  large  ungui- 
culate  dermal  pedicellariae.  In  this  character  and  in  general  appear- 


SHALLOW-WATER   STARFISHES  47 

ance  it  resembles  Coronaster,  but  the  latter  has  only  two  rows  of 
ambulacral  feet.  I  would  suggest  Heterasterias,  as  a  genus;  type, 
H.  volsellata. 

Genus  Marthasterias  Jullien. 

Marthasterias  JULLIEN,  Bull.  Zool.  Soc.  France,  p.  141,  1878. 
Stolasterias  (pars)  SLADEN,  Voy.  Challenger,  xxx,  p.  563,  1889.    Perrier,  Voy. 
Trav.  et  Talism.,  pp.  108,  109;  Resultats  Camp.  Scient.,  I,  p.  34,  1896. 

Rays  five,  angular,  normally  with  three  dorsal  radial  rows,  besides 
the  superomarginal  rows,  of  stout  plates  bearing  large,  mostly  conical 
spines ;  usually  two  on  the  superomarginals.  Inferomarginal  plates 
usually  with  two  rows  of  spines.  Adambulacral  plates  generally 
monacanthid,  each  plate  with  a  single  spine.  Peractinal  ossicles 
small  and  without  spines,  scarcely  visible  externally.1  When  young 
it  has  only  one  row  of  dorsal  spines,  the  median,  developed;  when 
very  large  there  may  sometimes  be  more  than  three  rows. 

Besides  the  type,  M.  gladalis  Miiller,  this  group  includes 
M.  africana,  of  South  Africa,  and  M.  rarispina  (Perrier). 

The  allied  diplacanthid  species  were  referred  to  another  generic 
group  (Distolasterias)  by  Perrier. 

Genus  Distolasterias  Perrier. 
Distolasterias  PERRIER,  Resultats  des  Camp.  Scient.,  i,  p.  34,  1896. 

This  name  was  proposed  by  Perrier  for  a  group  of  which  Stol- 
asterias stichantha  Sladen,  of  Japan,  was  the  type.  It  is  distin- 
guished from  his  Stolasterias = Marthasterias  by  the  diplacanthid 
arrangement  of  the  adambulacral  spines ;  and  by  having  five  or  more 
regular  close  longitudinal  rows  of  dorsal  plates.  The  actinal  plates 
are  rudimentary  and  without  spines. 

But  the  Stolasterias  neglecta  of  Perrier  (1896,  p.  37,  pi.  i,  figs.  2, 
za,  4-4^)  is  diplacanthid  and  therefore  does  not  go  in  that  genus,  as 
he  defines  it,  for  in  the  same  work  (op.  cit.,  p.  35)  he  makes  the 
genus  monacanthid.  In  this  respect  neglecta  is  like  his  Distolasterias 
(op.  cit.,  p.  34),  but  the  type  of  the  latter  has  a  much  more  complex 
dorsal  skeleton,  with  five  or  more  rows  of  dorsal  plates  and  spines. 
5".  neglecta  apparently  belongs  with  a  group  of  which  A.  forreri  Lor. 
may  be  taken  as  the  type. 

1  For  description  and  figures  of  skeleton,  see  Viguier,  op.  cit.,  pp.  100-105, 
pi.  v,  figs,  i-io,  1878. 


48  VERRILL 

Genus  Orthasterias  Verrill,  nov. 

Rays  long,  usually  five,  bearing  three,  five,  or  more  dorsal  rows 
of  rather  long  spines,  besides  a  superomarginal  row  on  each  side. 

Inferomarginals  bear  each  one  or  two  spines,  usually  two.  One 
row  of  interactinal  or  peractinal  plates,  with  or  without  spines,  close 
to  the  adambulacrals ;  the  latter  are  diplacanthid ;  a  short  subactinal 
row  may  also  occur  rarely. 

Lateral  and  dorsal  dermal  pedicellarise  are  large,  partly  wedge- 
shaped  or  spatulate,  often  dentate,  or  unguiculate.  Type,  O.  colum- 
biana  Ver.,  sp.  nov.  Also  includes  O.  dawsoni  Ver.,  nov. ;  O.  tanneri 
Ver. ;  O.  biordinata  Ver. ;  O.  calif ornica  Ver.,  etc. 

Subgenus  Stylasterias  Verrill,  nov. 
Type,  5".  forreri  Lor. 

Dorsal  plates  in  the  adult  usually  in  three  regular  rows,  having 
elongated  spines.  Upper  and  lower  marginal  plates  large  and 
regular,  with  a  well  marked  channel  between,  crossed  by  the  large 
descending  apophyses  of  the  upper  plates,  and  carrying  papulae  and 
large  forficiform  pedicellariae.  Lower  marginal  plates  close  to  the 
adambulacrals,  each  bearing  one  or  two  long  spines.  Peractinal 
plates  lacking  or  rudimentary;  when  present  never  bearing  spines, 
and  so  small  as  to  be  easily  overlooked,  without  dissection.  Adam- 
bulacral  spines  diplacanthid. 

Among  other  species  that  belong  to  this  group  are  the  following : 

S.  enopla  Ver.,  off  Nova  Scotia.1 
5".  neglecta  (Per.),  East  Atlantic. 
5".  contorta  (Per.),  West  Indies. 

Genus  Cosmasterias  Sladen,  1889. 

This  name  was  proposed  for  the  group  having  as  the  first  species, 
Asterias  (Cosmasterias)  tomidata  Sladen  (op.  cit.,  pi.  cv,  figs.  8-10), 
and  including  also  Asterias  sulcifera  Perrier,  1869  =  C.  lurida  (Phil- 
ippi),  1858. 

Perrier  has  adopted  Cosmasterias  with  C.  sulcifera1  (Per.),  from 
off  Patagonia,  etc.,  as  the  type. 

The  genus  is  diplacanthid  and  has  several  regular  rows  of  inter- 
actinal spines,  but  it  has  large,  unguiculate  or  felipedal,  dermal, 

1  For  full  synonymy  and  new  descriptions  of  this  species  and  C.  tomidata, 
see  Leipoldt,  Asterioidea  der  Vettor  Pisani  Exped.,  1895,  pp.  552-563.  See,  also, 
below,  under  Geographical  Distribution. 


SHALLOW-WATER   STARFISHES  49 

dorsal  pedicellariae,  not  found  in  northern  polyactinoplacid  types.  Its 
principal  dorsal  plates  are  in  regular  radial  rows. 

This  suggests  an  independent  line  of  development,  and  hence  the 
group  may  well  receive  a  distinctive  name.  C.  tomidata  (Sla.),  the 
type,  was  from  Chile,  in  45  fathoms. 

Somewhat  similar,  large,  unguiculate  or  felipedal  pedicellariae 
occur  on  Stichaster  polygrammus  Sla.,  off  Patagonia ;  S.  felipes  Sla., 
South  Africa;  Stolasterias  volsellata  Sla.,  Philippines;  Coronaster 
briar eus  (V.),  Atlantic,  and  some  other  species.1  In  these  they  are 
not  so  sessile  and  rigidly  erect  as  in  Pisaster.  Each  of  those  species 
named  is  now  referred  to  a  distinct  genus. 

E.  TABLE  OF  PRINCIPAL  GENERA  OF  ASTERIIN^E  WITH  REGU- 
LAR LONGITUDINAL  ROWS  OF  DORSAL  PLATES  AND 
SPINES. 

I.    NOT    MORE   THAN    ONE    ROW    OF   EVIDENT   ACTINAL    (PERACTINAL) 
PLATES, — MONACTINOPLACID  OR  ANACTINOPLACID. 

A.  Monacanthld,  or  nearly  so. 

B.  Rays  slender,  variable,  mostly  six  to  eleven;  often  more  than  two  madre- 

poric  plates  when  adult;  usually  autotomous. 

C.  One  (often  incomplete)   row  of  actinal  plates,  closely  united  to  adambu- 

lacrals,  and  usually  bearing  a  simple  row  of  spines   (monactino- 
placid).     Major  pedicellariae  large,  compressed,  ovate  or  pointed. 
Usually  three  or  five  rows  of  dorsal  plates  and  spines, 
a.  Peractinal  spines  usually  present  in  the  adult. 

Coscinasterias  Ver.     Type,  C.  muricata  Ver. 
aa.  Peractinal  spines  mostly  absent. 

Section  Stolasterias  (Sla.),  restr.    Type,  C.  (S.)  tenuispina. 
CC.  No  visible  actinal  plates,  the  inferomarginals  joining  the  adambulacrals 

(anactinoplacid).     Only   one   row   of   dorsal   plates — the   median   '-. 
radials.    Some  large,  unguiculate,  dermal  major  pedicellariae;  some 
of  the  plates  may  be  diplacanthid. 
Heterasterias  Ver.,  nov.    Type,  H.  volsellata  (Sla.) 

BB.  Rays  five  or  six,  stout.  Three  or  five  rows  of  strong  dorsal  plates  and 
spines.  No  interactinal  plates  with  spines.  Usually  but  one  madre- 
poric  plate ;  rarely  two. 

Marthasterias  Jullien.     Type,  M.  glacialis  (Mull.). 
AA.  Adambulacral  plates  mostly  diplacanthid. 

D.  No  evident  interactinal  plates  that  bear  spines;  if  present,  rudimentary. 

a.  Five  or  more  rows  of  stout  dorsal  plates  and  spines  when  adult.     Major 

pedicellariae  not  felipedal. 
Distolasterias  Per.     Type,  D.  stichantha  (Sla.). 

1  They  often  resemble,  more  or  less,  a  pair  of  miniature  clasped  hands  with 
fingers  interlocked  or  two  catspaws  with  claws  protruded  and  interlocking, 
and  hence  they  may  be  called  felipedal.  The  name  of  5".  felipes  Sla.  refers 
to  this  feature. 


50  VERRILL 

aa.  Only  three  regular  rows  of  dorsal  plates  with  spines. 

Stylasterias  Ver.,  nov.    Type,  S.  forreri  (Lor.), 
aaa.  Usually  only  one  row  of  dorsal  plates  (median)   developed;  they  are 

broad,  covered  with  a  thick  channeled  dermis. 
Sclerasterias  Per.    Type,  S.  guernei  Per.    E.  Atlantic. 
DD.  One  row  of  interactinal  plates  without  or  with  spines  (monactinoplacid). 
Three  to  five  or  more  rows  of  dorsal  plates  and  spines.     Major 
pedicellariae  of  two  kinds.    Some  compressed,  lanceolate  or  ovate; 
others  large,  thick,  spatulate  or  wedge-shaped.    Rays  five  or  six. 
Orthasterias  Ver.,  nov.    Type,  O.  columbiana  Ver.,  nov. 

II.  TWO  OR  MORE  ROWS  OF  ACTINAL  PLATES  AND  SPINES 
(POLYACTINOPLACID)  . 

A.  Diplacanthid.     Special  large  dermal  pedicellariae  are  unguiculate  or  feli- 

pedal. 
a.  Spines  vesiculated. 

Cosmasterias  Sla.    C.  tomidata  (Sla.). 
aa.  Spines  not  vesiculated. 

Cosmasterias  lurida  (Phil.)  —sulcifera  Per. 

F.  SUBDIVISIONS  OF  ASTERIIN^  WITH  RETICULATED  DORSAL 

SKELETONS. 

This  large  group  presents  fewer  morphological  characters  for  gen- 
eric and  subgeneric  distinctions  than  the  last,  and  some  of  them 
present  gradations  and  intermediate  conditions.  However,  if  we 
separate  the  genus  Pisaster  and  consider  the  large  group  to  which 
Asterias  rubens,  A.  vulgaris,  and  A.  forbesi  belong  as  typical 
Asterias,  there  will  be  left  several  more  or  less  differentiated  types  of 
structure. 

Genus  Leptasterias  Verrill. 
Type,  L.  mulleri  (Sars.). 

This  group,  as  separated  by  me,  1866,  differs  from  true  Asterias 
plainly  in  having  only  a  few  large  papulae  in  each  cluster.  Its 
typical  species  are  L.  mulleri,  L.  compta,  L.  tenera,  etc.  In  these 
the  dorsal  spines  are  slender  and  numerous  and  the  interactinal  spines 
are  in  one  row  (rarely  two).  The  discovery  of  numerous  additional 
species,  of  larger  size,  and  having  more  papulae,  now  renders  it  diffi- 
cult to  make  any  sharp  distinction,  in  this  respect,  between  the  two 
groups.  Hence  I  have  sometimes  considered  it  as  of  only  subgeneric 
value,  and  closely  connected  with  typical  Asterias.  Its  typical  species 
carry  the  large  eggs  and  young  attached  to  the  oral  region,  and  the 
development  is  direct  or  abbreviated. 


SHALLOW- WATER   STARFISHES  51 

The  most  important  generic  feature  is  the  fact  that  the  genital 
pores  are  on  the  actinal  side,1  between  the  proximal  inferomarginal 
plates,  not  dorsal,  as  in  Asterias.  The  ovaries  consist  of  compara- 
tively few  large  digitate  or  glomerate  tubules  which,  when  filled 
with  the  large  ova,  appear  beaded.  The  genital  ducts  are  short  and 
wide. 

Genus  Evasterias  Verrill,  nov. 
Type,  E,  troschelii  (Stimpson). 

We  may  also  distinguish  this  as  a  generic  group,  typified  by 
A.  troschelii.  In  this  the  dorsal  plates  and  ossicles  are  small  and 
reticulated,  while  there  are  three  or  more  regular  rows  of  stout  inter- 
actinal  spines  and  plates.  The  increased  number  of  rows  of  inter- 
actinal  plates  and  their  closer  union  are  the  principal  distinctions  in 
this  case,  but  the  number  of  plates  varies  with  age,  so  that  the  young 
specimens  are  essentially  like  Asterias  in  this  respect. 

Genus  Urasterias  Verrill. 

Plate  LXX,  figures  1-4. 
Type,  U.  linckii  (Muller  and  Troschel.)  =  A.  stellionura  Perrier. 

This  type  differs  so  much  in  structure  from  all  others  named  that 
it  should  probably  be  considered  a  distinct  genus. 

The  dorsal  skeleton  is  very  feebly  developed,  consisting  mostly  of 
small  plates  and  delicate,  short,  rod-like  or  linear  ossicles,  united  in 
an  irregular  network,  with  large  meshes,  but  having  a  median 
radial  row  of  rather  larger,  slightly  lobed,  spiniferous  plates,  and 
some  similar  lobed  plates  at  the  intersections  of  many  of  the  meshes. 
The  dorsal  spines  are  slender,  irregularly  scattered,  except  the 
radials.  Disk  covered  with  a  network  of  stouter,  somewhat  lobed 
ossicles,  bearing  solitary  spines,  and  usually  having  in  the  type  a 
larger  flat  plate,  or  group  of  such  plates,  at  each  interradial  area, 
above,  and  extending  to  the  lower  side,  and  usually  perforated  by  a 
'pair  of  papular  areas.  This  peculiar  group  of  plates  seems  to  be 
made  up  of  a  considerable  number  of  plates  in  the  adult,  often  united 

1  This  is  true  also  of  the  genus  Sporasterias  of  southern  South  America  and 
the  Antarctic  Ocean,  including  5".  antarctica  (Liitk.),  the  type  =  A sterias 
rugispina  Stiinp.  =  A.  spirabilis  Bell  =  A.  varia  (Philippi)  ;  S.  rupicola  Ver.; 
and  several  other  nominal  species;  and  probably,  also,  of  A.  studeri  Per.; 
Anasterias  chirophora  Lud. ;  A.  belgicce  Lud. ;  A.  perrieri  (Stud.)  ;  Podasterias 
lutkeni  (Per.)  ;  Stichaster  nutrix  (Stud.),  which  carry  their  young  in  the  same 
or  a  similar  manner.  They  are  all  antarctic. 


52  VERRILL 

together  in  young  specimens.  Upper  and  lower  marginal  plates 
wide  and  strong,  each  bearing,  in  the  young,  a  single  spine.  The 
upper  marginal  plates  have  a  large  and  wide  descending  lobe ;  these 
join  the  stout  upper  lobe  of  the  inferomarginals,  but  leave  wide 
papular  areas  between  them,  along  the  wide  lateral  channels.  The 
upper  marginals  have  small  side  lobes,  which  are  united  together 
longitudinally,  by  smaller  narrow,  interpolated  spineless  plates. 

Peractinal  plates  are  small,  or  rudimentary  without  spines.  The 
adambulacrals  generally  appear  to  be  joined  directly  to  the  infero- 
marginals, but  minute  ossicles  may  intervene.  Adambulacral  plates 
diplacanthid  or  subdiplacanthid.  (PI.  LXX,  fig.  I.) 

Major  pedicellariae  are  large  and  numerous,  especially  between  the 
inferomarginal  spines  and  on  the  inner  margins  of  the  grooves,  but 
also  occur  on  the  back.  They  are  acute-ovate  and  a  little  com- 
pressed. The  minor  pedicellariae  are  unusually  large  and  denticu- 
late; they  form  very  large  pedicellate  clusters,  especially  on  the 
inferomarginal  spines.  Ambulacra!  pores  and  feet  very  large. 

The  lack  of  distinct  peractinal  plates  and  spines  and  the  presence  of 
the  large  interradial  plates  are  very  important  morphological  char- 
acters, aside  from  the  feeble  dorsal  skeleton.  The  large  size  of  the 
ambulacral  pores  is  a  notable  feature. 

The  type,  U.  linckii  =  stellionura  (Per.),  is  found  from  Spitz- 
bergen  to  Nova  Scotia.1  Another  arctic  form,  U.  gunneri  (Dub.  and 
Kor.),  is  closely  allied  and  may  be  only  a  variety  (teste  Ludwig). 
One  or  both  of  these  may  very  likely  be  found  hereafter  in  Bering 
Sea,  for  they  are  probably  circumpolar. 

Another  arctic  species,  U.  panopla  (Stuxberg)  seems  to  be  con- 
generic with  U.  linckii.  It  has  a  similar,  very  feeble,  reticulated  dorsal 
skeleton,  and  lacks  distinct  peractinal  plates  and  spines.  Its  proximal 
adambulacral  plates  in  some  cases  bear  three  or  four  spines,  but  are 
mostly  diplacanthid.  The  superomarginal  and  inferomarginal  plates 
are  cruciform  and  are  connected  by  a  narrow,  spineless,  intermediate 
plate.  The  inferomarginal  ossicles  usually  bear  two  spines,  some- 
times more  in  large  examples.  It  is  from  Spitzbergen,  Barents  Sea, 
Kara  Sea,  Finmark,  etc.  It  is  remarkable  for  the  great  number 
and  large  size  of  the  dorsal  minor  pedicellariae. 

1It  has  been  found  as  a  fossil  abundant  in  the  postglacial  clays  at  Salem, 
Massachusetts. 


SHALLOW-WATER   STARFISHES  53 

Genus  Parasterias  Verrill,  nov. 
Type,  P.  albertensis  Verrill,  British  Columbia. 

Five  rays,  stout;  dorsal  plates  openly  reticulated,  much  as  in 
Asterias;  interactinal  spines  and  plates  lacking ;  adambulacrals  dipla- 
canthid.  For  more  details  see  below,  p.  187. 

Genus  Ctenasterias  Verrill,  nov. 
Type,  C.  spitzbergensis  (Dan.  and  Koren). 

Dorsal  skeleton  composed  of  an  irregular  network  of  slender,  but 
numerous,  mostly  transverse  ossicles,  bearing  large  numbers  of 
minute,  uniform  spines,  arranged  in  transverse  rows  or  combs  on 
the  sides  of  the  rays.  Marginal  spines  in  two  double  rows  near 
together.  Adambulacral  plates  diplacanthid,  or  partly  triplacanthid. 
Two  forms  of  major  pedicellariae ;  the  larger  ones  are  lyrate  or 
cylindrical  and  obtuse.  A  single  row  of  small  interactinal  spines. 
Probably  closely  allied  to  Leptasterias.  Reproduction  not  known. 

C.  cribaria  (Stimpson)  is  pretty  certainly  identical  with  the  type, 
which  is  from  Spitzbergen. 

Genus  Allasterias  Verrill. 
Type,  A.  rathbuni  (Verrill).    Bering  Sea. 

Adambulacral  spines  alternately  diplacanthid  and  triplacanthid,  or 
monacanthid  and  diplacanthid;  the  inner  spine  of  the  larger  alter- 
nate group  is  inserted  on  an  angle  of  the  plate  within  the  edge  of  the 
furrow.  Dorsal  skeleton  openly  reticulated;  superomarginal  plates 
and  spines  form  an  angular  margin. 

Genus  Sporasterias  Perrier. 

Type,  5".  antarctica  (Liitk.)  =A.  rugispina  Stimp.  (teste  Leipoldt). 
Adambulacral  spines  regularly  monacanthid.  Dorsal  skeleton 
irregularly  reticulated,  sometimes  nearly  as  in  some  species  of  Lept- 
asterias, sometimes  weakly  developed.1  Genital  pores  ventral;  the 
eggs  and  young  are  carried  over  and  around  the  mouth,  as  in  Lept- 
asterias. For  details,  see  below,  under  Geographical  Distribution,  p. 
355- 

Genus  Smilasterias  Sladen. 
Type,  S.  scalprifera  (Sladen).    Off  Patagonia,  etc. 
Adambulacral  plates  diplacanthid  or  triplacanthid,  the  spines  and 
those  of  the  inferomarginal  plates  large,  flat,  and  placed  in  transverse 
rows.    Dorsal  skeleton  irregularly  reticulated  with  small  spines. 

*In  5".  rupicola  Ver.,  from  Kerguelen  Island,  the  dorsal  skeleton  is  well 
developed,  with  short,  stout  ossicles  and  numerous  small  spines. 


54  VERRILL 

Genus  Meyenaster  Verrill. 
Type,  Af.  gelatinosus  (Meyen,  1834).     Chile. 
Meyenaster  VERRILL,  op.  cit,  1913,  p.  485. 

Adambulacral  plates  monacanthid ;  one  row  of  larger  interactinal 
spines  on  thick  ovate  plates,  each  in  line  with  two  similar  spines  on 
each  inferomarginal  plate;  superomarginal  plates  stout,  each  alter- 
nate one  with  a  single  large  spine ;  a  wide  intermarginal  lane  bearing 
a  regular  row  of  papular  areas,  twice  as  many  as  superomarginal 
spines,  and  many  large  pedicellariae.  A  row  of  median  radial  stout 
spines,  usually  one  on  each  alternate  plate;  in  large  specimens  an 
intermediate  imperfect  row  on  each  side;  medio-lateral  skeleton 
reticulated  irregularly  with  numerous  short,  transverse  and  oblique 
ossicles  defining  five  or  six  irregular  series  of  large  papular  areas 
on  each  side.  Central  part  of  disk  symmetrically  stellate  with  a 
central  and  twelve  surrounding  spines,  in  type.  Large,  stout  pedi- 
cellariae dorsal,  lateral,  and  intra-ambulacral,  compressed,  ovate-tri- 
angular, not  unguiculate ;  many  very  small  acute,  major  pedicellariae 
on  papular  areas ;  minor  pedicellariae  abundant  on  large,  broad,  fleshy 
circumspinal  sheaths.  A  thick,  strongly  canaliculated  skin  covers  and 
conceals  all  the  plates. 

G.  ANALYTICAL  TABLE  OF  THE  GENERIC  AND  SUBGENERIC 
GROUPS  OF  NORTHERN  ASTERIIN^  HAVING  RETICULATED 
DORSAL  SKELETONS. 

A.  Monacanthid.     Dorsal   skeleton   coarsely   reticulate.     Interactinal   spines 

and  ossicles  well  developed,  in  several  rows.    Large  unguiculate 
erect  pedicellariae  are  present;  rays  five  or  six. 
Pisaster  (Ag.).    Type,  P.  ochraceus  (Brandt). 

AA.  Usually  diplacanthid,  or  alternately  diplacanthid  and  monacanthid,  rarely 
triplacanthid.  Interactinal  plates  and  spines  present  or  absent,  or 
rudimentary. 

B.  Adambulacral  spines  confined  to  the  outer  or  actinal  surface  of  the  plate. 

C.  Dorsal  plates  mostly  angular  or  lobed.     Interactinal  plates  and  spine* 

forming  at  least  one  row  (peractinal),  usually  with  other  shorter 
proximal  rows.    Both  rows  of  marginals  well  developed. 
Asterias,  sens  ext. 

D.  Dorsolateral  plates  and  transverse  ossicles  openly  reticulated. 

a.  Interactinal  plates  strong,  closely  united,  or  imbricated  in  three  or  more 
rows  in  the  adult,  bearing  several  regular  close  rows  of  nearly 
uniform  spines,  like  the  inf eromarginals ;  intervening  papular  areas 
small.  Two  kinds  of  major  pedicellariae  usually  present  Dorsal 
spines  unequal,  several  on  a  plate,  reticulate  or  areolate. 
Genus  Evasterias  V.  Type,  E.  troschelii  (St.). 


SHALLOW-WATER   STARFISHES  55 

aa.  Interactinal  plates  not  so  closely  united;  usually  one  or  two  rows,  with 
larger  papular  areas  between.  Major  pedicellariae  usually  of  but  one 
kind.  Dorsal  spines  rather  irregularly  arranged,  usually  scattered, 
sometimes  areolate  or  acervate. 

b.  Papular  areas  large,  bearing  numerous  small  papulae.    Larvae  are  usually 

free-swimming  brachiolariae ;  genital  pores  dorsal. 
Asterias,  restr.    Type,  A.  rubens  L. 

bb.  Papular  areas  smaller  and  bearing  fewer  large  papulae.    Eggs  and  larvae 
are  usually  carried  by  the  mother  around  the  mouth.     No  free 
larval  stages ;  genital  pores  ventral. 
Genus,  Leptasterias  Ver.    Type,  L.  mulleri  (Sars). 

aaa.  Interactinal  plates  lacking  or  rudimentary;  papular  areas  large,  numer- 
ous. 

Parasterias,  nov.    Type,  P.  albertensis  Ver.,  nov. 

DD.  Dorsolateral  plates  and  ossicles  wide,  irregular,  rather  closely  joined, 
leaving  only  small  papular  areas  in  longitudinal  lines,  but  the 
ossicles  do  not  form  regular  rows.  All  the  dorsal  and  marginal 
plates  are  covered  with  close  clusters  of  very  small  spines,  longer 
on  the  inferomarginal  and  interactinal  plates.  Autotomous;  rays 
mostly  two  or  three  to  nine;  finally  six  in  adult. 
Stephanasterias  Ver.  Type,  S.  albula  (Stimp.).  Circumpolar;  North 

Atlantic;  Bering  Sea. 

CC.  Dorsal  plates  mostly  rod-like.  Interactinal  plates  small  or  rudimentary, 
mostly  spineless.  Usually  two  kinds  of  major  pedicellariae. 

c.  Dorsal  skeleton  feeble,  composed  mostly  of  small,  linear  or  roundish,  iso- 

lated and  reticulated  ossicles.  Dorsal  spines  few  and  mostly  scat- 
tered, but  usually  forming  a  median  radial  row.  Both  rows  of  mar- 
ginal plates  well  developed  and  rather  large,  bearing  single  or 
double  rows  of  stout  spines,  separated  by  a  wide  lateral  channel. 
Dermal  major  pedicellarise  large  and  abundant,  wedge-shaped. 
Minor  pedicellariae  abundant  on  the  spines  and  skin,  attached  to  a 
large  fleshy  sheath  on  the  spines.  Alternate  superomarginals  with- 
out spines. 

Urasterias  Ver.    Type,  U.  linckii  (Mull,  and  Trosch.). 

cc.  Dorsal  skeleton  composed  of  nearly  uniform  short  linear  rods,  irregularly 
reticulated,  and  covered  with  numerous  small,  uniform  spinules, 
often  in  transverse  rows  or  combs  on  the  sides.    Two  rows  of 
marginal  plates  near  together,  in  the  type  bearing  double  rows  of 
spines.    Larger  major  pedicellarise  obtuse,  large. 
Ctenasterias  Ver.  nov.    Type,  C.  spitzbergensis  (D.  and  KO- 
BE. The  inner  adambulacral  spine,  on  alternate  plates,  is  attached  to  the  inner 
edge  of  the  plate,  or  within  the  groove.     Peractinal  plates  feebly 
developed  or  lacking.    Dorsal  spines  small  and  numerous. 
Allasterias  Ver.    Type,  A.  rathbuni  V. 

H.  ASTERIID^E  OF  THE  NORTHWESTERN  COAST  OF  AMERICA. 
This  faunal  region  contains  a  remarkable  assemblage  of  the  repre- 
sentatives of  this  family.  A  few  species  from  north  of  the  Aleutian 
Islands  are  of  arctic  or  circumpolar  origin,  but  most  are  endemic. 
The  most  remarkable  genus  is  Pycnopodia,  strictly  confined  to  this 


56  VERRILL 

region.  The  genus  Pisaster  is  here  represented  by  seven  or  eight 
large  species,  some  of  them  being  among  the  largest  and  most  mas- 
sive starfishes  known.  It  is  doubtful  if  the  restricted  genus  occurs  in 
any  other  fauna.  The  genus  Orthasterias,  with  seven  or  eight 
species,  is  uncommon  elsewhere.  Its  nearest  allies  are  Atlantic 
species  and  Distolasterias  of  Japan. 

A  large  proportion  (fifteen)  of  the  species  of  this  region  are 
normally  six-rayed,  which  is  a  remarkable  peculiarity. 

Over  twenty  nominal  species  of  Asteriinae  had  been  already  de- 
scribed from  the  Northwest  Coast,  between  southern  California  and 
the  Arctic  Ocean.  In  the  collections  studied  by  me  there  are  about 
thirty  additional  species,  besides  twenty  well-marked  new  varietal 
forms,  or  a  total  of  about  seventy. 

Probably  no  other  fauna  is  so  rich  in  species  of  this  group.1  It 
is  not  improbable,  however,  that  some  of  the  supposed  species  may 
prove  to  be  only  local  varieties,  when  larger  series  can  be  carefully 
studied.  This  may  possibly  prove  true  of  some  of  the  larger  forms 
described  by  Stimpson,  allied  to  Pisaster  ochraceus  and  P.  capitatus, 
but  on  the  other  hand  there  are  probably  many  additional  species  to 
be  found  by  dredging  in  moderate  depths.  Nearly  all  the  known 
species  of  this  region  are  littoral  or  very  shallow-water  forms. 
Deep-water  species  are  not  described  in  this  report. 

I.  ANALYTICAL  TABLE  OF  THE  GENERA,  SUBGENERA,  SPECIES, 
SUBSPECIES  AND  VARIETIES  OF  ASTERIIDJE  FROM  THE 
NORTHWEST  COAST  OF  AMERICA. 

1.  Dorsal  skeleton  continuous,  formed  mostly  of  lobed  plates  and  smaller 

ossicles  overlapping  at  the  ends;  dorsal  plates  not  isolated.    Rays 
five  or  six,  rarely  more,  unless  autotomous. 

2.  Adambulacral  spines  all  attached  to  the  actinal  face  of  the  plates;  none 

attached  higher  within  the  grooves. 

3.  Adambulacral  spines  in  a  single  regular  row,  one  to  a  plate,  in  the  adult 

(monacanthid). 

A.  Interactinal  plates  present,  at  least  in  adults,  and  usually  bearing  spines. 

C.  Disk  thick,  rather  large ;  rays  five  or  six,  stout,  thick  at  base.  Dorsal  os- 
sicles strong,  numerous,  not  confined  to  three  or  five  rows.  Dorsal 
skeleton  reticulate  or  irregular.  Dorsal  and  lateral  dermal  major 
pedicellariae  sessile,  large,  stout,  erect,  ovoid  or  wedge-shaped,  often 
nearly  as  thick  as  the  spines,  and  usually  strongly  denticulated  at 
the  tips  of  the  valves  (unguiculate). 
Genus  Pisaster  M.  and  Tr.,  p.  67.  Type,  P.  ochraceus  (Br.). 

*The  only  other  region  that  can  be  compared  with  this  in  the  number  and 
variety  of  Asteriinae  is  the  southern  coast  of  South  America,  especially  around 
Tierra  del  Fuego  and  Patagonia,  whence  numerous  genera  and  species  have 
been  described,  mostly  very  unlike  the  northern  forms.  (See  below,  p.  351.) 


SHALLOW-WATER   STARFISHES  57 

D.  Interactinal  ossicles  numerous,  stout,  closely  united  in  three  to  six  longi- 

tudinal rows,  usually  bearing  three  to  six  or  more  close  rows  of 
strong  actinal  spines. 

E.  Dorsal  spines  arranged  in  a  reticulate  or  areolate  manner,  or  else  grouped 

on  the  nodes ;  but  often  forming  a  median  row  on  the  rays.  Rays 
normally  five,  stout. 

a.  Dorsal  spines  stout,  coarse,  very  unequal,  capitate,  acervate,  forming  con- 

spicuous angular  reticulations  with  or  without  notable  nodular 
groups  and  often  with  median  radial  rows. 

J^  P.  ochraceus  (Brandt),  p.  68.    Calif,  to  Middle  Alaska, 

a'.  Dorsal  spines  mostly  in  very  prominent  nodes  or  clusters ;  reticulations  less 

conspicuous. 

Var.  nodiferus  V.,  nov.,  p.  71.    Calif,  to  Alaska. 

aa.  Dorsal  spines  small,  or  rather  small,  acute  or  obtuse,  not  strongly  capitate, 
usually  forming  evident  reticulations. 

b.  Dorsal  spines  exceedingly  numerous,  very  small,  obtuse  or  acute,  forming 

conspicuous  reticulations,  with  or  without  radial  rows  and  a  cen- 
tral pentagon.  Actinal  spines  stout,  obtuse,  but  not  strongly  grooved 
externally. 

P.  confer tus  (Stimp.),  p.  73.    Calif,  to  Vancouver  I. 

bb.  Dorsal  spines  small,  slender,  not  very  numerous,  forming  an  openly  reticu- 
lated pattern  with  a  central  pentagon  and  distinct  median  radial 
rows.  Actinal  spines  deeply  gouge-shaped  and  often  bifid.  Disk 
large. 

P.  fissipinus  (Stimp.),  p.  76.    Oregon. 

EE.  Dorsal  spines  scattered,  mostly  without  regular  order,  not  reticulated, 
but  sometimes  forming  radial  rows.  Rays  usually  stout,  five  or  six. 

d.  Rays  normally  five.    Dorsal  spines  large,  with  capitate  or  ovoid  tips,  not 

very  numerous. 

e.  Dorsal  spines   few,  large,  nearly  equal,  obtuse  or  strongly  capitate  and 

radially  striate.    Rays  short. 
P.  capitatus  (Stimp.),  p.  81.     S.  California. 

ee.  Dorsal  spines  more  numerous,  unequal,  the  tips  ovoid  or  subconic  and 
striated,  usually  forming  imperfect  radial  rows.  Rays  long,  rounded. 
Size  large. 

P.  lutkenii  (Stimp.),  p.  83.    S.  Calif,  to  Vancouver  I.  Var.  australis 
Ver.,  p.  88. 

dd.  Rays  normally  six;  dorsal  spines  very  numerous,  short,  stout,  conical, 
acute  or  obtuse,  striated,  nearly  uniformly  scattered ;  no  evident  ra- 
dial rows.     Interactinal  spines  in  four  to  six  rows.     Disk  large. 
Size  large. 
P.  giganteus   (Stimp.),  p.  89. 

DD.  Interactinal  ossicles  fewer,  usually  forming  only  two  rows,  and  mostly 
bearing  simple  rows  of  spines.  Dorsal  ossicles  openly  reticulated. 
Dorsal  spines  few. 

f.  Rays  normally  five. 

g.  Dorsal  spines  rather  sparse,  short,  thick,  subconic,  striated,  scattered  or 

clustered,  with  distinct  median  radial  rows.    Actinal  spines  short, 
obtuse,  often  flat,  or  slightly  gouge-shaped  at  tip. 
P.  brevispinus  (Stimp.),  p.  77.     S.  Calif,  to  Vancouver  I. 


58  VERRILL 

gg.  Rays  thick  and  high  at  base,  rapidly  tapered.  Dorsal  spines  few,  stout, 
blunt,  or  conical,  mostly  isolated,  but  forming  imperfect  median 
radial  rows  and  a  regular  pentagon  on  the  disk.  Superomarginal 
spines  one  to  a  plate.  Inferomarginal  row  double.  Large  dermal 
pedicellariae  numerous,  stout,  stone-hammer-shaped,  with  serrate 
jaws. 
P.  papulosus  Ver.,  p.  91.  Puget  Sd.,  Brit.  Columbia. 

ff.  Rays  normally  six,  rarely  five;  more  slender.    Dorsal  spines  short,  rather 
numerous,  obtuse  or  capitate,  irregularly  arranged,  often  in  groups 
of  two  or  three.    Interactinal  spines  in  two  or  three  rows.    Major 
pedicellariae  acute-triangular. 
P.  grayi  Ver.  =  katherinae  Per.  (non  Gray),  p.  97.    Oregon. 

CC.  Rays  five;  angular,  more  slender.  Dorsal  spines  in  three  or  five  regular 
rows.  Dorsal  ossicles  stout  in  longitudinal  and  transverse  rows. 

h.  One  or  two  rows  of  interactinal  plates  and  spines.     Major  pedicellariae 
large,  ovate;  minor  pedicellariae  not  very  abundant.     Superomar- 
ginal spines  one  to  a  plate. 
P.  paucispinus  (Stimp.),  p.  98.     Calif,  to  Puget  Sd. 

AA.  Dorsal  spines  in  three  regular  rows,  bearing  dense  subapical  clusters  of 

minor  pedicellariae.    Superomarginal  spines  only  on  alternate  plates. 

Ventral  spines  in  two  or  three  rows.     Interactinal  plates  feebly 

developed. 

Marthasterias  sertulifera  (Xantus),  p.  100.    Gulf  of  Calif,  to  S.  Diego. 

3'.  Adambulacral  spines  two  to  a  plate,  arranged  in  two  rows  (diplacanthid)  ; 
or  else  in  irregular  rows,  when  some  of  the  plates  have  one  and 
some  two  spines  (subdiplacanthid). 

B.  At  least  one  row  of  visible  interactinal  plates  generally  present  in  adults, 
often  spineless;  sometimes  several  close  rows.  -  *£  *& 

F.  Dorsolateral  plates  rather  small,  narrow,  strongly  lobed,  usually  reticulated 

or  irregularly  arranged,  and  united  by  their  own  lobes  and  by 
smaller  transverse  ossicles. 

G.  Interactinal  plates  usually  form  one  long  row  (peractinal),  much  like  the 

lower  marginals,  and  one  or  two  short  proximal  rows  (subactinals), 
but  the  latter  may  be  mostly  lacking  or  rudimentary,  especially  in 
the  young,  so  that  there  may  be  only  two  simple  rows  of  ventral 
spines ;  or  the  lower  marginals  and  peractinal  rows  may  be  doubled 
proximally,  two  or  more  spines  standing  on  each  plate;  thus  the 
number  of  actual  rows  of  these  spines  may  sometimes  be  four  or 
more.  Dorsal  ossicles  variously  arranged,  most  often  openly  reticu- 
late. Major  pedicellariae  usually  small,  ovate  or  lanceolate,  some- 
times unguiculate.  Rays  normally  five  or  six. 

H.  Dorsal  papular  areas  usually  large;  papulae  small  and  numerous,  in  groups 
of  twelve  to  fifty  or  more,  in  the  adult.    Size  generally  rather  large. 
Larva  of  many,  if  not  all,  species  is  a  free-swimming  brachiolaria. 
Genital  pores  dorsal. 
Genus  Asterias  (L.),  closely  restricted,  p.  101. 

i.  Rays  normally  five,  ossicles  and  spines  reticulate  in  arrangement. 


SHALLOW-WATER   STARFISHES  59 

j.  Rays  five,  rather  long,  rounded.  Primary  dorsal  spines  few,  short,  stout 
(2  to  3  mm.  thick),  capitate,  forming  a  median  row,  with  few 
irregularly  scattered  ones  of  similar  size  and  form.  Two  marginal 
and  two  interactinal,  regular,  mostly  simple,  rows  of  short,  stout, 
blunt  spines;  adambulacrals  subdiplacanthid. 
A.  victoriana  Ver.,  p.  102.  Vancouver  I. 

jj.  Rays  long,  slender,  acute;  dorsal  spines  of  moderate  length,  slender, 
fluted,  mostly  one  to  a  plate,  arranged  in  quincunx,  or  in  about 
seven  obscure  alternating  rows;  median  row  with  longer  spines, 
distinct;  both  marginal  rows  single;  peractinal  row  double,  with 
spines  longer  than  marginals;  major  pedicellariae  adambulacral, 
rather  large,  lanceolate. 
A.  namimensis  Ver.,  p.  105.  Brit.  Columbia. 

ii.  Rays  normally  six. 

k.  Dorsal  spines  very  diverse  in  form  and  size;  primary  ones  very  large,  ir- 
regularly placed,  but  not  acervate,  very  stout  (2  to  4  mm.),  with 
nipple-shaped,  sulcated  tips;  smaller  spines  numerous,  unequal, 
capitate  or  round-topped,  with  abundant  minor  pedicellariae.  Mar- 
ginal and  interactinal  spines  smaller,  in  about  four  close  rows 
proximally;  adambulacrals  mostly  two  to  a  plate;  the  inner  ones 
and  furrow  margins  bear  abundant  clusters  of  small  major  pedicel- 
lariae. 
A.  polythela  Ver.,  p.  104.  Arctic  O.  ~f-t  **f,  7°,  ?*•  •  * V 

kk.  Primary  dorsal  spines  not  unusually  large;  not  nipple-shaped. 

I.  Primary  dorsal  spines  more  or  less  unequal,  capitate;  the  larger  usually 

acervate  or  irregularly  grouped.  Pedicellariae  small,  numerous; 
major  ones  ovate,  forming  clusters  on  the  adambulacral  spines,  a 
few  also  on  the  interactinal  areas.  One  complete  row  of  peractinal 
spines,  separated  from  the  adambulacrals  by  a  naked  zone  contain- 
ing papular  areas.  Subactinal  spines  form  a  short  series,  or  may  be 
lacking. 
A.  acervata  Stimp.,  p.  107.  Bering  Sea;  Arctic  O.  ft'  Z7i  *ob 

II.  Principal  dorsal  spines  not  very  unequal  nor  distinctly  acervate. 

m.  Dorsal  spines  rather  small,  irregularly  reticulated;   adambulacrals   sub- 
diplacanthid. 
A.  katherina  Gray  (non  Per.),  p.  112.    Oregon;  Gulf  of  Georgia. 

mm.  Dorsal  spines  very  numerous,  in  distinst  radial  rows,  not  very  diverse  in 
size,  capitate,  several  on  a  plate;  the  median  row  crowded  and 
more  or  less  clustered ;  both  marginal  rows  double ;  two  interactinal 
rows. 
A.  multiclava  Ver.,  p.  114.    Siberia.      fd. 

HH.  Papular  areas  small;  papulae  few,  large,  mostly  standing  singly  or  in 
small  groups  of  one  to  five,  rarely  ten  or  more.  Dorsal  plates 
usually  lobed  and  united  by  overlapping,  sometimes  in  radial  rows. 
Adambulacral  spines  alternately  one  or  two  to  a  plate,  or  else 
biserial.  Minor  pedicellariae  as  in  Asterias;  large,  serrate,  dermal 
ones  may  occur.  Eggs  and  young  usually  carried  in  clusters  around 
the  mouth  of  the  mother;  genital  pores  ventral.  Size  small.  Rays 
normally  five  or  six. 
Genus  Leptasterias  Ver.,  in  part,  p.  116.  (See  below,  p.  60.) 


6O  VERRILL 

I.  Rays  normally  five,  sometimes  varying  individually  to  six. 

n.  Disk  somewhat  enlarged,  thickened  or  depressed;  rays  not  very  slender; 
spines  unequal  and  uneven;  superomarginal  row  mostly  single; 
inferomarginal  row  double. 

o.  Disk  and  rays  depressed;  dorsal  spines  very  unequal;  larger  ones  capitate, 
forming  a   prominent  median   radial   row   and   irregularly   linear 
groups;  ventral  spines  all  rather  slender. 
L.  inequalis  Ver.,  sp.  nov.,  p.  117.    Orca,  Alaska. 

oo.  Disk  thickened  or  plump;  rays  strongly  convex;  dorsal  spines  numerous, 
capitate  or  clavate,  usually  areolated,  or  arranged  partly  in  clusters, 
often  forming  a  median  row ;  marginal  and  peractinal  spines  thick- 
ened. 
L.  epichlora  (Br.),  p.  132  (five-rayed  form).    Aleutian  Is.  to  Puget  Sd. 

nn.  Disk  smaller,  rays  more  slender  or  terete.  Dorsal  spines  subequal,  clus- 
tered, scattered,  or  somewhat  in  longitudinal  rows. 

p.  Dorsal  spines  not  remarkably  small;  not  in  transverse  lateral  combs.  At 
least  one  row  of  interactinal  spines  proximally. 

q.  Rays  very  slender,  terete;  dorsal  spines  small,  capitate,  or  clavate,  numer- 
ous, mostly  one  or  two  to  a  plate,  not  in  evident  rows.     Ventral 
spines  slender,  both  marginal  and  peractinal  rows  single;  major 
pedicellariae  large,  with  long  spatulate  blades. 
L.  leptalea  Ver.,  sp.  nov.,  p.  119.    Virgin  Bay,  Alaska. 

qq.  Rays  less  slender.    Dorsal  spines  small,  clavate,  in  about  three  longitudi- 
nal rows,  not  crowded;  both  marginal  rows  regular  and  distinct, 
the  spines  longer  and  standing  singly;  interactinal  row  imperfect. 
L.  arctica  (Murdoch),  p.  120.    Arctic  O. ;  Bering  Sea, 

pp.  Dorsal  spines  very  slender,  numerous  irregularly  arranged ;  superomarginal 
and  inferomarginal  spines  longer,  mostly  one  to  a  plate,  in  regu- 
lar rows. 

-^•x  L.  epichlora  miliaris  Ver.,  nov.,  p.   138  (five-rayed  form).     British 
Columbia. 

ppp.  Dorsal  and  marginal  spines  very  numerous,  small,  subequal,  arranged  in 
close  clusters,  and  largely  concealed  by  the  large  dense  wreaths  of 
minor  pedicellariae;  marginal  plates  of  both  series  have  two  or 
three  small  spines,  nearly  concealed  by  pedicellariae. 
Leptasterias  obtecta  Ver.,  sp.  nov.,  p.  144.    Bering  Sea. 

nnn.  Dorsal  and  dorso-marginal  spines  very  small,  numerous,  nearly  equal, 
the  latter  arranged  in  transverse  lines  on  the  narrow  ossicles  defin- 
ing large  transverse  papular  areas.  Minor  pedicellariae  small, 
around  bases  of  spines,  not  unusually  abundant. 

Ctenasterias  cribraria   (Stimp.),  p.   148.     Circumpolar;  Bering  Sea; 
Arctic  O. ;  North  Atlantic. 

II.  Rays  normally  six,  sometimes  five  or  seven ;  not  autotomous. 

r.  Disk  small;  rays  slender,  not  enlarged  at  base;  papulae  few,  single  or  in 
small  groups.    Ambulacral  pores  of  the  usual  size  and  form,  more  or 
less  crowded,  in  four  rows. 
Genus  Leptasterias  Ver.,  in  part,  p.  116.     (See  above,  p.  59.) 

t.  Dorsal  spines  small,  slender,  clavate,  not  crowded  nor  clustered,  pretty 
regularly  arranged  in  quincunx,  surrounded  by  thick  wreaths  of 


SHALLOW-WATER   STARFISHES  6l 

pedicellarise ;    marginal   spines   longer;    rows   mostly   simple;   no 
peractinal  spines. 
L.  coei  Ver.,  sp.  nov.,  p.  123.    Off  Berg  Bay,  Alaska. 

tt.  Dorsal  spines  longer,  scattered;  one  or  two  to  a  plate,  except  in  evident 
median  radial  row.     Inferomarginal  row  double,  two  to  a  plate, 
longer  than  dorsals,  subacute;  superomarginal  row  simple;  three 
interactinal  rows  proximally,  one  spine  to  a  plate. 
L.  macouni  Ver.,  sp.  nov.,  p.  124.     Vancouver  I.  >-*  -fa 

ttt.  Dorsal  spines  mostly  clustered,  several  on  each  plate. 

u.  Rays  six  or  seven,  elongated.  Dorsal  spines  small,  obtuse,  arranged  in 
five  longitudinal  ranges  of  groups,  each  group  with  three  or  four 
spines ;  others  are  on  the  transverse  ossicles,  so  that  they  form  "  a 
vague  reticulation."  Ventral  spines  in  two  rows,  pointed,  longer 
than  adambulacrals. 
L.  vancouveri  (Per.),  p.  125.  Vancouver  I.  ^t*V*f- 

uu.  Dorsal  spines  small  and  numerous,  all  of  about  the  same  height,  usually 
clavate,  obtuse,  standing  in  crowded  rows.  Marginal  and  inter- 
actinal spines  notably  longer,  forming  three  to  five  or  more  close 
rows. 

v.  Dorsal  spines  small,  slender,  subequal,  mostly  clavate,  not  much  crowded, 
standing  singly  or  in  small  groups  on  the  ossicles,  forming  median 
radial  rows,  and  sometimes  subradial  series.     Major  pedicellariae 
few,  rather  large,  long,  ovate. 
L.  hexactis  (St.),  p.  126.     Calif,  to  Brit.  Columbia.      V/Jr, 

w.  Dorsal  spines  mostly  capitate,  very  small  and  very  numerous,  standing  in 
large  groups  on  each  plate,  about  equal  in  height,  and  thus  pre- 
senting a  nearly  even  surface.  Papular  pores  in  nearly  regular 
radial  rows.  Marginal  and  interactinal  spines  much  longer,  often 
bent,  numerous,  arranged  in  regular  rows.  Peroral  spines  strong. 
L.  aqualis  (St.),  p.  128.  Calif,  to  Puget  Sd. 

w.  Dorsal  spines  very  short,  equal,  clavate  or  capitate,  crowded,  many  on  each         *  ~ 
plate.     Marginal   and  interactinal   spines   form  about   four  rows. 
Rays  short,  obtuse. 
Var.  nana  Ver.,  nov.,  p.  132.    California;  Oregon;  British  Columbia. 

ww.  Dorsal   spines   very  numerous   and   crowded  on   the  plates,   distinctly 
capitate,  with  rounded  tips.    Marginal  and  interactinal  spines  more 
numerous,  forming  five  to  eight  rows,  several  on  each  plate. 
Var.  compacta  Ver.,  nov.,  p.  130.     California. 

www.  Dorsal  spines  very  numerous,  crowded,  the  clusters  arranged  in  three 
or  five  radial  bands  separated  by  rows  of  very  distinct  papular  areas. 
Var.  concinna  Ver.,  nov.,  p.   132.     California. 

rr.  Rays  normally  six,  sometimes  five,  of  moderate  length,  thick  or  wide  at 
base,  not  slender,  rather  obtuse;  disk  somewhat  wide  or  swollen, 
skeleton  firm;  principal  dorsal  ossicles  rather  large. 

x.  Dorsal  ossicles  not  very  closely  united,  irregularly  reticulated,  covered  with 
numerous  short,  mostly  capitate  or  clavate  spines,  usually  reticulate 
in  arrangement,  often  showing  a  more  or  less  evident  median  row 
or  band ;  three  rows  of  interactinal  spines  often  present  proximally 
in  adult,  only  one  in  small  specimens.  Adambulacral  spines  irregu- 


62  VERRILL 

larly   diplacanthid,   crowded;   sucker- feet  numerous,   crowded   in 
four  rows.     Large,  denticulate,  dermal  major  pedicellariae  often 
present  in  adult.     Eggs  and  young  carried  by  mother.     Papulae 
often  in  dorsal  clusters  of  five  to  twelve  or  more. 
L.  epichlora   (Br.),  p.   132.     Calif,  to  Aleutian  Is. 

a.  Rays  commonly  five;  dorsal  spines  very  numerous,  very  unequal,  strongly 

capitate,  acervate.    Median  row  not  very  evident 
Var.  subnodulosa  Ver.,  nov.,  p.  139.    Wrangel,  Alaska. 
aa.  Rays  normally  six. 

b.  Superomarginal  spines  stand  two  or  three,  or  sometimes  more,  to  a  plate. 

c.  Dorsal  spines  quite  unequal  in  size,  mostly  capitate;  arrangement  mostly 

areolated  or  reticulated. 

d.  Dorsal  spines  do  not  form  a  very  evident  median  row,  and  are  not  notably 

acervate;  reticulate  arrangement  conspicuous;  larger  ones  capitate. 
Large  denticulate  dermal  pedicellariae  often  present,  as  well  as  the 
usual  form ;  minor  sort  abundant 

Subspecies  alaskensis  Ver.,  nov.,  p.  136.     Puget  Sd.  to  Aleutian  Is. 
dd.  Dorsal  spines  form  an  evident  median  row  or  band  of  larger  spines. 

e.  Dorsal  spines  numerous,  mostly  clavate,  not  crowded,  reticulate,  with  the 

median  radial  row  very  evident. 
Var.  carinella  Ver.,  nov.,  p.  137.    Alaska. 

ee.  Median   radial   dorsal   spines   decidedly   larger,    capitate,    others    small, 

crowded,  reticulate  and  areolate. 
Var.  siderea  Ver.,  nov.,  p.  137.    Puget  Sd.  to  Aleutian  Is. 

eee.  Sometimes  five-rayed.    Dorsal  spines  crowded,  very  small,  slender,  cla- 
vate or  slightly  capitate;  several  small  ones  around  each  larger; 
median  ones  slightly  larger;  superomarginals  unequal,  one  larger 
and  several  smaller  on  a  plate. 
Subspecies  miliaris  Ver.,  nov.,  p.  138.    Alaska. 

cc.  Dorsal    spines,    mostly    equal    or    subequal,    short,    capitate,    numerous, 
crowded,  areolate,  often  forming  circles  around  the  papular  areas. 

1  Dorsal  spines  are  in  five  or  more  evident  radial  bands  separated  by  rows  of 
large  papular  areas,  several  spines  standing  on  each  of  the  larger 
ossicles,  median  band  evident,  but  not  prominent;  three  to  four  or 
more  spines  on  each  superomarginal  plate;  two  on  inferomarginals, 
stouter;  one  on  peractinal,  stout,  clavate.  Papular  areas  large.  No 
large  pedicellariae  on  type. 
Subspecies  plena  Ver.,  nov.,  p.  140.  British  Columbia. 

ff .  Dorsal  spines  very  even,  stout,  capitate,  crowdedly  areolate,  but  not  forming 
evident  radial  rows ;  median  row  irregular  or  obscure ;  papular  areas 
conspicuous,  but  not  in  evident  radial  rows;  mostly  three  spines 
on  superomarginal  plates;  two  on  inferomarginals,  all  similar  and 
near  together,  clavate,  larger  and  longer  than  dorsals;  peractinals 
similar.  Large,  erect,  serrate  major  pedicellariae  common  between 
marginal  rows  of  spines. 
Subspecies  pugetana  Ver.,  nov.,  p.  142.  Puget  Sd. 

bb.  Superomarginal  spines  mostly  stand  singly  on  the  plates.     A  distinct 
dorsal  median  row. 


SHALLOW-WATER   STARFISHES  63 

g.  Dorsal  spines  small,  subequal,  slender,  clavate. 

Var.  regularis  Ver.,  nov.,  p.  139.    Alaska. 
gg.  Dorsal  spines  short,  mammilliform,  arranged  somewhat  in  rows. 

Var.  subregularis  Ver.,  nov.,  p.  139.    Alaska. 

xx.  Dorsal  and  marginal  ossicles  stout,  closely  united,  overlapping;  median 
dorsal  radial  row  prominent,  bearing  the  larger  capitate  spines  in  a 
conspicuous  irregular  row;  other  spines  of  the  larger  sort  form 
an  irregular  radial  row  on  each  side;  inferomarginal  spines  longer 
and  larger,  two  or  three  to  a  plate;  single  row  of  peractinals  simi- 
lar. Disk  is  rather  wide,  depressed. 
L.  dispar  Ver.,  nov.,  p.  142.  Aleutian  Is. 

GG.  Interactinal  and  submarginal  plates  rather  stout,  lobate,  closely  joined  in 
three  to  five  regular  longitudinal  rows;  each  plate  may  bear  one, 
two  or  more  spines,  forming,  in  the  adult,  four  to  eight  or  more 
rows  of  uniform  or  similar  ventral  spines.  Rays  five,  rounded, 
usually  long  and  tapered.  Dorsal  ossicles  mostly  reticulate.  Papu- 
lar areas  large.  Unguiculate  dorsal  pedicellariae  lacking. 
Genus  Evasterias  Ver.,  nov.,  p.  151. 

L  Rays  five,  gradually  tapered.    Dorsal  spines  numerous,  unequal,  areolate 
or  reticulate,  often  clustered;  larger  ones  short,  capitate  or  trun- 
cate; smaller  ones  acute,  capitate  or  clavate.    Marginal  spines  in 
regular  rows;   longer  than  the  dorsals;   interactinal  spines  still 
longer,  forming  four  to  six  or  more  regular  close  rows,  curving 
strongly  upward  proximally;  adoral  spines  elongated. 
E.  troschelii  (St.),  p.  151.    Calif,  to  Alaska. 
j.  Dorsal  spines  acervate,  or  forming  very  evident  nodular  groups  with  larger 

central  capitate  spines,  and  often,  also,  a  median  row. 
Var.  subnodosa  Ver.,  nov.,  p.  163.    Puget  Sd.  to  Alaska, 
jj.  Dorsal  spines  not  acervate. 

k.  Dorsal  spines  smaller,  very  numerous,  mostly  clavate  or  subacute,  few 
capitate,  rough,  openly  reticulate;  papular  areas  large;  numerous 
crowded  interactinal  spines.    Size  large. 
Var.  rudis  Ver.,  nov.,  p.  158.    Vancouver  I.,  etc. 
kk.  Dorsal  spines  nearly  all  capitate,  not  very  unequal. 
1.  Dorsal   spines   nearly   uniform,   capitate,   forming   close   reticulations,   the 

spines  in  single  lines  on  the  ossicles. 
Var.  dcnsa  Ver.,  nov.,  p.  161.    Vancouver  I. 

1L  Rays  of  moderate  length,  dorsal  spines  all  capitate,  reticulations  coarse; 
both  marginal  rows  of  spines   simple  or  nearly  so,   spines  not 
crowded;  two  or  three  actinal  rows,  not  crowded. 
Var.  alveolata  Ver.,  nov.,  p.  162.    Vancouver  I.  to  Alaska, 
kkk.  Dorsal  spines  not  very  unequal,  nearly  all  slender,  clavate  or  subclavate, 
areolate,  not  acervate;  marginals  larger,  subcapitate.    Pedicellarise 
numerous. 

Var.  parvispina  Ver.,  nov.,  p.  163.     Alaska. 

ii.  Rays  five,  long  and  large,  acute  at  tips.  Dorsal  spines  very  numerous, 
small,  unequal,  the  larger  obtuse.  The  smaller  ones  are  mostly 
acute  and  arranged  in  clusters  and  short  transverse  lines  or  combs, 
producing  an  imperfectly  reticulate  pattern,  without  distinct  radial 


64  VERRILL 

rows.    Interactinal  and  submarginal  spines  form  five  or  six,  mostly 
double,  crowded  rows,  with  about  eight  to  ten  in  each  transverse 
range,  in  the  adult.    Proximal  adambulacral  spines  much  elongated. 
Size  large. 
E.  acanthostoma  Ver.,  nov.,  p.  165.    Alaska. 

FF.  The  dorso-lateral  spines,  or  groups  of  spinules,  and  more  or  less  of  the 
plates,  have  an  evident  arrangement  in  longitudinal  rows.  Inter- 
actinal plates  present  or  rarely  absent;  seldom  more  than  one  row. 

J.  Dorsal  plates  strong,  lobate,  reticulated  so  as  to  form  rather  obvious 
longitudinal  rows;  papular  areas  usually  large,  often  nearly  rec- 
tangular. Each  of  the  principal  plates  usually  bears  a  single  rather 
large  spine,  rarely  two,  forming  three  to  five  or  more  dorsal  rows 
of  spines;  smaller  spines  may  occur  on  the  intermediate  transverse 
ossicles.  Marginal  plates  stout,  bearing  regular  rows  of  spines, 
much  like  the  dorsals.  A  row  of  similar  peractinal  plates,  usually 
bearing  a  simple  row  of  spines,  often  spineless,  rarely  lacking ;  some- 
times a  short  row  of  subactinals  proximally.  Adambulacrals  usually 
diplacanthid,  sometimes  subdiplacanthid.  Major  pedicellariae  large; 
usually  of  two  or  more  kinds ;  the  larger  dermal  ones  denticulate,  or 
unguiculate.  Rays  five  or  six. 

Genus   Orthasterias  Ver.,  nov.,  p.   168.     Type,   O.  columbiana  Ver., 
sp.  nov. 

K.  A  row  of  interactinal  plates  is  present,  with  or  without  spines;  they  may 
be  small  and  not  visible  unless  dermis  be  removed.  TM;  "Jb  (P  •» 

L.  One  row  of  peractinal  spines,  at  least  proximally;  inferomarginals  with 
two  rows,  making  three  ventral  rows  altogether.—  t»  l»fe 

a.  Rays  normally  five,  rarely  six,  elongated. 

b.  Dorsal  spines  in  at  least  five  rows,  besides  some  intermediate  scattered 

ones  on  the  connective  ossicles ;  papular  areas  rather  small ;  dorsal 
radial  plates  rather  large  and  thick,  firmly  united. 

c.  Five  dorsal  radial  rows  of  spines  obvious. 

d.  Apical  oral  spines  not  notably  enlarged.    Dorsal  spines  obtuse  and  fluted. 

Wreaths  of  minor  pedicellariae  on  thick  dermal-sheaths,  often  above 
the  middle  of  the  spine. 

e.  Adambulacral  spines   flattened;   adoral   spines   longer,  not   very  slender; 

dorsal  spines  obtuse  or  truncate,  rough  or  fluted. 

O.   columbiana   Ver.,   sp.   nov.,   p.    168.     Vancouver   I.   to  Yakutat, 

Alaska. 

ee.  Adambulacral  spines  all  very  slender  and  subequal;  regularly  diplacan- 
thid, strongly  and  evenly  divergent,  thus  forming  two  remarkably 
regular  pectinate  rows,  those  of  the  inner  row  usually  horizontal. 

O.  biordinata  Ver.,  nov.,  p.  173.    British  Columbia. 

dd.  Apical  oral  spines  notably  stronger  than  others.  Dorsal  spines  tapered, 
not  fluted.  Minor  pedicellariae  are  notably  minute,  forming  dense 
basal  circumspinal  wreaths.  Major  pedicellariae  are  very  large, 
stout,  often  as  thick  as  spines,  wedge-shaped,  erect,  often  dentic- 
ulate. 

O.  californica  Ver.,  nov.,  p.  174.     California. 


SHALLOW-WATER   STARFISHES  65 

cc.  Longitudinal  rows  of  spines  more  than  five,  not  very  obvious,  due  to 
crowding ;  spines  appear  as  if  in  quincunx. 

f.  Dorsal    spines    long,    slender,    slightly    grooved.       Adambulacral    spines 

slender,  orals  longer.    Major  pedicellariae  large,  lanceolate,  acute, 
dentate  at  tip. 

O.  kcehleri  Lor.,  p.  175.    Vancouver  I. 

bb.  Dorsal  spines  in  three  rows,  long,  not  fluted;  ossicles  openly  reticulated, 
leaving  large  papular  areas ;  spines  of  two  inferomarginal  rows  and 
peractinal  row  slender,  elongated,  tapered;  adambulacral  spines 
rather  slender,  tapered  or  slightly  flattened.  Major  pedicellariae 
large. 

0.  dawsoni  Ver.,  nov.,  p.  175.    Brit.  Columbia. 

aa.  Rays  six,  rounded  and  elongated.  Dorsal  spines  large,  in  three  or  five 
rows,  with  smaller  ones  interpolated,  all  with  a  thick  fleshy  sheath 
(in  alcohol)  bearing  a  large  wreath  of  minor  pedicellariae  at  about 
mid-height  or  above.  Both  marginal  rows  of  spines  are  regular 
and  mostly  simple.  A  regular  peractinal  row  of  spines  and  in 
mature  specimens  a  short  subactinal  row  proximally. 

O.  merriami  Ver.,  nov.,  p.  177.    Alaska. 

LL.  Peractinal  spines  generally  lacking;  the  ossicles  are  rudimentary  or  small; 
inferomarginals  with  two  rows  of  spines. 

Subgenus  Stylasterias  Ver.,  nov.,  p.  179. 

g.  Dorsal  spines  mostly  in  three  longitudinal  rows,  often  with  some  scattered 

intermediate  ones. 

h.  Minor  pedicellarise  remarkably  large  and  unguiculate,  about  half  as  long 
as  major  ones,  forming  large,  loose,  basal  circumspinal  wreaths. 
Larger  major  pedicellariae  often  nearly  as  stout  as  the  spines. 

i.  Spines  long,  inferomarginals  in  two  rows  and,  like  the  outer  adambulacral 
spines,  often  notably  flattened  and  spatulate  or  gouge-shaped.  Outer 
adambulacral  spines  are  the  larger;  papular  areas  large;  peractinal 
plates  very  small. 
O.  forreri  (Lor.),  p.  179.    California. 

ii.  Adambulacral  plates  and  spines  crowded;  spines  slender.    Minor  pedicel- 
lariae remarkably  large  and  abundant;  dorsal  ones  in  large  groups      ^  l^vv 
on  the  dermis  and  around  bases  of  spines;  dorso-lateral  ossicles       S^L*v" 
rather  slender,  openly  reticulated;  papular  areas  large;  major  pedi- 
cellariae very  stout. 
Subspecies  forcipptlata  Ver.,  p.  180.    British  Columbia. 

hh.  Minor  pedicellariae  not  notably  large ;  they  form  large,  dense,  basal  circum- 
spinal wreaths;  inferomarginal  and  adambulacral  spines  flattened, 
not  notably  spatulate  nor  strongly  gouge-shaped.  Dorsal  spines 
well  spaced,  elongated,  tapered,  not  fluted ;  alternate  supermarginal 
plates  mostly  spineless. 
O.  gonolena  Ver.,  nov.,  p.  184.  S.  California  to  Gulf  of  Calif. 

gg.  Dorsal  spines  in  more  than  three  rows;  rays  slender,  terete. 

Dorsal  spines  in  about  five  rows,  long,  nearly  uniform,  tapered; 
papular  areas  large;  rays  slender;  inferomarginals  tapered,  obtuse; 
adambulacrals  small,  slender,  outer  ones  a  little  larger  and  longer, 
slightly  clavate.  Major  pedicellariae  large,  denticulate. 


66  VERRILL 

O.  leptolena  Ver.,  nov.,  p.  182.    British  Columbia. 
"'WC.  No  visible  interactinal  plates  or  spines.    (See  below,  p.  66.) 
Genus  Distolasterias  Per.,  in  part,  p.  185. 

Dorsal  spines  numerous,  in  about  seven  to  nine  irregular  alter- 
nating rows;  rather  short,  fluted,  blunt  or  clavate.  Larger  major 
pedicellariae  partly  stout,  wedge-shaped;  partly  elongated,  spatulate, 
unguiculate,  with  few  large  curved  teeth.  Adambulacral  spines 
slender,  crowded,  regularly  diplacanthid. 

_., D.  chelifera  Ver.,  nov.,  p.  185.    Vancouver  I. 

JJ.  Dorso-lateral  plates  mostly  wide,  lobed,  overlapping  or  imbricated  in  few 
close  rows  (stichasterial  arrangement),  covered  with  clusters  of 
minute  spinules.  A  distinct  peractinal  row  of  plates.  Disk  small, 
rays  slender,  six  in  type.  Ambulacral  pores  unusually  large,  sub- 
triangular  or  cordate,  not  crowded.  Dorsal  spines  very  small, 
short,  scattered,  obtuse  or  clavate,  a  distinct  median  radial  series. 
Stenasterias  tnacropora  Ver.,  p.  145.  Alaska. 

BB.  Interactinal  plates  lacking  or  rudimentary ;  no  interactinal  spines. 
M.  Dorsal  skeleton  weak,  composed  mostly  of  slender,  openly  reticulated 
ossicles,  some  of  which  do  not  unite  in  a  continuous  skeleton.  Dor- 
sal spines  few,  rather  large,  in  few  open  rows  or  scattered;  upper 
marginal  spines  regular  and  well  separated  from  lower  marginals; 
the  latter  with  large  and  long  spines ;  alternate  plates  without  spines. 
Minor  pedicellariae  large  and  very  numerous,  in  sacculated  wreaths 
or  clusters.  Size  large. 

Genus  Urasterias  Ver.,  p.  187.    Type,  U.  linckii  (M.  and  T.).    Arctic; 
North  Atlantic. 

b.  Several  open  rows  of  long  dorsal  spines ;  adambulacral  plates  not  crowded ; 

lower  marginal  spines  large  and  well  spaced;  minor  pedicellariae 
not  remarkably  large;  bunched  in  large  wreaths  on  the  large,  loose, 
saccular  sheaths  of  the  spines.  Size  large. 

U.  linckii  (M.  and  T.),  p.  187  (extralimital?).    N.  Atlantic  and  Arctic. 
MM.  Dorsal  skeleton  well  developed. 

c.  Dorso-lateral  plates  and  ossicles  openly  reticulated.     Rays  regularly  five, 

rather  short,  stout,  depressed;  dorsal  spines  numerous,  small, 
slender,  obtuse,  arranged  on  small,  slender,  reticulated  ossicles;  no 
distinct  median  row;  marginal  spines  usually  two  to  a  plate  in  both 
rows,  in  form  like  the  dorsals,  but  rather  longer;  adambulacrals 
regularly  diplacanthid.  No  interactinals  visible. 
Genus  Parasterias  nov.,  p.  187.  Type,  P.  albertensis  Ver.  Brit. 

Columbia. 

cc.  Dorso-lateral  spines  and  plates  in  more  or  less  obvious  longitudinal  or 
transverse  rows ;  spines  either  solitary  or  clustered. 

d.  Rays  regularly  five,  large  and  long;  dorsal  plates  bear  large  and  mostly 

solitary  spines  and  large  pedicellariae. 

Genus  Distolasterias  Per.,  in  part,  p.  185.    (See  above,  p.  65.) 

dd.  Rays  slender,  variable  in  number,  due  to  autotomy;  disk  small;  autoto- 

mous;  finally  six  or  five.    Median  and  marginal  plates  little  wider, 

imbricated  in  longitudinal  rows ;  dorso-lateral  ossicles  and  plates 

short,  irregularly  reticulated,  but  leaving  regular  rows  of  papular 


SHALLOW-WATER  STARFISHES  67 

areas.    All  the  plates  are  covered  with  close  clusters  of  small,  nearly 
equal  spinules,  longer  beneath.    Pedicellariae  of  two  sorts;  dermal, 
not  abundant ;  few  on  spines. 
Stephanasterias  albula  (St.),  p.  147.     Circumpolar;  North  Atlantic; 

Bering  Sea. 

2'  The  inner  adambulacral  spine,  on  alternate  plates,  is  attached  to  a  lobe  of 
the  inner  edge  of  the  plate,  higher  up  within  the  groove.    Dorsal 
spines  small  and  numerous,  scattered.     Interactinal  plates  feebly 
developed  or  lacking. 
Allasterias  Ver.,  p.  188.    Type.  A.  rathbuni  Ver.    Bering  Sea. 

f.  Dorsal  ossicles  small,  forming  reticulations ;  adambulacral  spines  alternately 

two  and  three  to  a  plate.  Superomarginal  spines  clustered.  Dermal 
major  pedicellariae  large  and  numerous,  pointed.  Rays  five,  wide, 
depressed;  margin  prominent. 

A.  rathbuni  Ver.,  p.  189.     Bering  Sea.     See  also,  A.  forficulosa  V., 
nov.,  p.  194.    Japan. 

g.  Inferomarginals  not  very  stout,  flattened,  blunt,  not  deeply  channeled. 

Subspecies  nortonensis  Ver.,  p.  191.    Bering  Sea.      K»ri»*.  (frt**M>  • 
gg.  Inferomarginal  and  actinal  spines  stout,  channeled  or  gouge-shaped. 

A.  anomala  Ver.,  p.  193.  Siberia.  (  T^t^t  -|v*m  3t  /V\«eV**4  X 
1'.  Dorsal  skeleton  incomplete  or  rudimentary;  ossicles  partly  isolated;  dorsal 
surface  covered  by  soft  skin.  Disk  large;  rays  multiple,  variable 
in  number,  increasing  with  age  by  interpolation  in  pairs  up  to  twenty 
to  twenty-four  in  the  adult.  Adambulacral  spines  monacanthid. 
Marginal  plates  distinct.  Dorsal  minor  pedicellariae  in  large  clus- 
ters, dermal  and  epispinal. 

Genus  Pycnopodia  Stimp.,  p.  197.    Type,  P.  helianthoides  (Br.),  p.  198. 
Calif,  to  Aleutian  Islands. 

ASTERIINyE :  DESCRIPTIONS  OF  GENERA  AND  SPECIES. 
Genus  Pisaster  A.  Agassiz,  ex  Miiller  and  Troschel. 

Pisoster  MULLER  and  TROSCHEL,  Syst.  Ast.,  p.  20,  note,   1842.     A.  Agassiz, 

North  American  Starfishes,  p.  96,  1877  (type,  ochraceus). 
Cosmasterias  (pars)  SLADEN,  Voy.  Challenger,  xxx,  p.  562,  1876! 

Large  starfishes  with  a  thick  and  usually  broad  disk  and  either 
five  or  six  stout  tapering  rays.  Adambulacral  spines  form  a  single 
regular  row,  one  to  each  plate  (monacanthid).  Three  to  six  rows 
of  closely  united  or  tesselated,  thick  actinal  and  lower  marginal 
plates,  bearing  several  close  rows  of  stout  ventral  spines.  The  syn- 
actinal  plates  are  closely  united  to  the  adambulacrals.  Oral  region 
usually  deeply  sunken.  Interbrachial  septa  well  developed. 

Special,  very  large,  stout,  erect,  sessile,  forficulate  pedicellariae, 
with  interlocking  serrations  at  the  ends  of  the  valves  (unguiculate), 
stand  on  the  lateral  channels,  and  often  between  the  actinal  and 
dorsal  spines ;  larger  ones  often  stand  in  the  interradial  areas  below. 

Dorsal  skeleton  variable,  but  well  developed,  the  ossicles  generally 


68  VERRILL 

stout  and  openly  reticulated.  Dorsal  spines  few  or  many,  and 
variously  arranged,  often  in  reticulate  or  acervate  patterns,  but 
sometimes  scattered  or  in  irregular  rows. 

One  of  the  important  structural  features,  characteristic  of  most 
of  the  species,  is  the  increased  number  of  rows  of  actinal  plates  and 
spines,  and  their  close  imbricated  or  tesselated  arrangement;  but 
in  certain  species  (P.  papulosus),  otherwise  related  to  the  type,  there 
are  usually  only  two  rows  of  actinal  plates  that  bear  spines  below  the 
inferomarginal  plates,  as  in  typical  Asterias;  very  large  specimens 
may  have  more  rows. 

But  there  is  another  character,  perhaps,  of  more  value,  common  to 
all  the  species  from  the  Northwestern  American  Coast  related  to 
P.  lutkenii  and  P.  ochraceus,  which  seems  to  warrant  the  separation 
of  this  group,  as  a  distinct  genus.  This  is  the  existence  of  the 
peculiar  form  of  large,  stout,  erect,  sessile,  dermal  pedicellariae, 
ovoid  or  wedge-shaped  in  form,  with  broad,  unguiculate  or  denticu- 
lated, interlocking  jaws.  These  occur  especially  along  the  naked 
spaces  between  the  ventral  and  lateral  spines,  but  they  are  also 
frequently  found  scattered  between  the  dorsal  spines,  or  between 
the  bases  of  the  actinal  spines.  They  are  often  equal  to  the  adjacent 
spines  in  diameter,  but  are  much  shorter.  More  slender,  lanceolate 
or  ovate,  major  pedicellariae,  of  the  ordinary  type,  always  coexist 
with  these  large  forms,  while  the  latter  are  not  present  on  starfishes 
of  the  Asterias  rub  ens  type,  nor  on  those  belonging  to  most  of  the 
other  divisions.1 

It  is  probable,  therefore,  that  this  group  had  its  origin  upon  the 
Pacific  coast  of  North  America,  where  it  has  subsequently  had  time 
enough  to  develop  into  many  diverse  species,  some  of  which  may 
have  migrated  to  other  regions  and  left  descendants.  This,  then,  is 
one  of  the  most  characteristic  groups  of  this  fauna,  but  it  is  not  so 
strikingly  peculiar  as  Pycnopodia  and  Dermasterias. 

Mr.  A.  Agassiz  (1877)  referred  to  the  name  Pisaster  as  having 
been  proposed  by  Professor  L.  Agassiz,  for  the  group  of  species 
allied  to  A.  ochracea,  but  he  did  not  consider  the  group  as  more  than 
a  subdivision  of  Asterias,  though  he  fully  described  the  characteristic 
structures  in  the  skeleton  of  P.  ochraceus,  as  contrasted  with  a 
typical  Asterias.  Miiller  and  Troschel  had,  however,  proposed  this 

*A  few  similar  large  pedicellariae  often  occur  on  Leptasterias  epichlora 
(Br.),  but  not  on  the  other  allied  species.  They  are  often  wanting,  and  may 
be  due  to  hybridizing  with  P.  ochraceus,  associated  with  it.  Similar  large 
ones  are  found  on  Orthasterias  columbiana  and  allied  species. 


>  .  SHALLOW- WATER   STARFISHES  69 

name,  for  the  same  type,  in  1842,  but  did  not  subsequently  use  it 
This  name  should,  therefore,  take  precedence  of  all  others,  whether 
it  be  considered  a  genus  or  a  subgenus.  • ' ,  "• 

The  mode  of  reproduction  is  unknown. 

PISASTER  OCHRACEUS  (Brandt)  A.  Agassiz. 
Plate  xxi,  figures  i,  2 ;  plate  XLIX,  figures  3-30?  (pedicellariae)  ; 

plate  LVI,  figures  3,  30  (variety). 

Aster  ms  ochracea  BRANDT,  Prod.  Descr.  Anim.  Mertens,  p.  69,  1835. 
Asterias  janthina  BRANDT,  op.  cit.,  p.  69,  1835  (descr.  insufficient;  color  var.  ?). 
Asterias  ochracea  STIMPSON,  Journ.  Boston  Soc.  Nat.  Hist.,  vi,  p.  87,  pi.  xxin, 
fig.  2,  1867.    Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  pp.  325,  326,  1867.    Whit- 
eaves,  Trans.  Royal  Soc.  Canada,  rv,  p.  116,  1887.     A.  Agassiz,  North 
Amer.   Starfishes,   Mem.   Mus.   Comp.   Zool.,  v,  p.  96,  pL   xi,  figs.   1-7 
(structure),  1877  (Pisaster).    H.  L.  Clark,  op.  cit.,  1907,  p.  67,  pi.  vi, 
fig.  3   (structure).     Loeb,  Publ.  Univ.  Calif.,  Physiol.,  n,  pp.  5-30,  1904 
(hybridization). 
Asteracanthion  margaritifer  MULLER  and  TROSCHEL,   Syst.,  p.  20,  1842   (t 

Bell). 

Disk  rather  large,  strongly  reticulated  by  the  prominent  ossicles 
and  spines ;  rays  normally  five,  sometimes  six,  stout,  rapidly  tapered. 
Two  Alaskan  specimens,  differing  considerably  in  form,  measure  as 
follows:  (a)  lesser  radius,  36  mm.;  greater  radius,  120  mm.;  ratio, 
1:3.33;  (b)  lesser  radius,  32  mm.;  greater  radius,  118  mm.;  ratio, 
i :  3.68. 

The  dorsal  spines  are  numerous,  unequal  in  size,  but  nearly  all 
are  capitate;  the  larger  ones  are  short  and  stout,  strongly  capitate 
with  flattened  striated  tops.  They  are  arranged  in  short  rows  form- 
ing a  conspicuously  reticulated  pattern,  with  large,  deep,  angular 
papular  areas  between  them.  They  usually  stand  so  closely  in  the 
rows  that  they  nearly  touch;  on  the  disk  they  form  a  conspicuous 
pentagon,  enclosing  subordinate  angular  groups  or  partial  reticula- 
tions and  a  central  cluster.  A  zigzag,  irregular  median  row  is  often 
distinct  on  the  basal  part  of  the  rays.  The  smaller  spines  are  similar 
in  form  but  not  so  stout ;  they  are  partly  grouped  around  the  larger 
ones,  and  partly  serve  to  subdivide  the  primary  reticulations.  Each 
primary  papular  area  usually  contains  several  large  clusters  of 
papulae.  Large,  short,  very  stout,  obtuse,  wedge-shaped,  unguiculate, 
dermal  major  pedicellariae  are  more  or  less  numerous  on  the  dorsal 
surface,  and  still  more  so  on  the  sides  of  the  rays;  they  are  rather 
less  in  diameter  than  the  adjacent  spines.  Those  in  the  lateral  chan- 
nels and  interradial  spaces  are  larger  and  stouter  than  the  dorsal 
ones.  Minor  pedicellariae  form  wreaths  around  the  bases  of  the 


7O  VERRILL 

dorsal  and  lateral  spines  and  also  occur  in  clusters  on  the  papular 
areas. 

The  interactinal  spines  form  four  or  five  crowded  rows ;  they  are 
large,  stout,  mostly  clavate,  with  obtuse,  smaller,  often  flattened, 
striated  or  sulcated  tips;  those  of  the  synactinal  (inner)  row  are 
less  stout,  but  not  shorter;  those  of  the  interradial  areas  are  longer 
and  more  fusiform.  Toward  the  mouth  the  adambulacral  rows  of 
ossicles  are  prolonged  as  a  narrow  ridge  to  the  sunken  mouth,  unac- 
companied by  actinal  spines.  The  actinal  spines  bear  clusters  of 
minute  minor  pedicellariae.  The  interactinal  ossicles,  as  exposed  in  a 
denuded  specimen,  are  small  and  closely  arranged  in  about  five  rows 
at  the  base  of  the  arm,  with  some  additional  ones  irregularly  inter- 
polated. Large,  pedicelled,  lanceolate  major  pedicellariae  are  attached 
within  the  margins  of  the  furrows.  The  marginal  spines  are  capi- 
tate and  striated  like  the  dorsal  spines. 

VARIATIONS. 

In  the  Museum  of  Comparative  Zoology  I  have  seen  two  six- 
rayed  specimens  (No.  1423)  from  Crescent  City,  Calif.  These  have 
the  dorsal  spines  unusually  numerous,  and  strongly  reticulated  or 
areolated. 

Two  specimens  sent  to  me  by  Mr.  J.  F.  Whiteaves,  of  the 
Canadian  Geological  Survey,  from  Vancouver  Island,  are  remarkable 
for  their  great  size.  They  are  20  to  22  inches  in  diameter.  Radii  of 
one  are  75  mm.  and  250  to  275  mm. ;  ratios  about  i :  3.75.  Radii  of 
the  other  are  70  mm.  and  240  to  260  mm. ;  ratios  about  1 :  3.6.  The 
disk  is  abnormally  flattened  in  both,  by  imperfect  preservation. 

They  agree  with  ordinary  specimens  in  all  essential  features, 
except  such  as  are  naturally  due  to  greater  size.  The  dorsal  spines 
are  unequal  and  strongly  clustered,  but  have  lost  more  or  less  of 
their  reticulated  arrangement,  except  distally,  owing  to  the  breaking 
up  of  many  of  the  lines  of  spines  and  the  interpolation  of  new 
clusters.  The  spines  and  pedicellariae  are  of  the  ordinary  forms. 
The  large,  serrate  pedicellariae  are  very  abundant  on  the  dorsal, 
lateral,  and  actinal  surfaces.  The  largest  are  on  the  actinal  inter- 
radial  areas,  where  some  equal  the  spines  in  diameter. 

Mr.  Whiteaves  also  sent  a  short-rayed  specimen,  from  the  Queen 
Charlotte  Islands,  which  has  the  dorsal  spines  larger,  more  capitate, 
and  more  numerous  than  usual,  so  that  they  form  very  prominent 
nodular  clusters  and  imperfect  reticulations.  Its  large  unguiculate 
pedicellariae  are  unusually  abundant.  (Var.  nodiferus.) 


SHALLOW-WATER   STARFISHES  Jl 

In  the  same  collection  was  a  large  six-rayed  specimen  from  Dis- 
covery Sound,  1885,  which  differs  in  no  other  respect  from  the 
ordinary  five-rayed  specimen.  A  similar  one,  in  our  collection,  was 
collected  near  Monterey,  Calif.,  by  Dr.  W.  R.  Coe. 

The  smallest  specimen  that  I  have  seen  has  the  radii  14  mm.  and 
41  mm. ;  ratio  about  1 : 2.8.  The  disk  is  elevated.  The  dorsal  sur- 
face is  thickly  covered  with  characteristic  nodular  groups  and  imper- 
fect reticulations  of  capitate  spines,  like  those  of  larger  specimens, 
but  smaller.  On  the  disk  they  form  a  distinct  stellate  pentagon  and  a 
central  cluster. 

The  upper  marginal  row  is  simple.  The  inferomarginal  row  is 
double.  There  are  also  two  simple  rows  of  actinal  spines ;  all  these 
forming  four  very  regular,  close-set  rows  of  nearly  equal  obtuse 
ventral  spines.  On  the  lateral  channels,  proximally,  there  are  a  few 
of  the  large,  erect,  wedge-shaped  pedicellariae,  characteristic  of  the 
large  specimens. 

Its  color  is  variable,  and  often  very  showy;  most  frequently  it  is 
dark  or  light  orange,  varying  to  yellow,  to  chocolate-brown,  and  to 
violet;  spines  are  paler. 

This  is  the  most  common  of  the  species  of  Pisaster,  from  Mon- 
terey and  San  Francisco  to  Sitka.  In  most  places  it  is  by  far  the 
most  abundant  large  starfish  of  the  shores.  It  occurs  at  low  tide 
among  rocks  and  in  shallow  water. 

It  was  collected  by  the  Harriman  Expedition  at  Sitka,  Virgin  Bay, 
Wrangel,  and  other  localities  in  southern  Alaska. 

Mr.  J.  F.  Whiteaves,  of  the  Canadian  Geological  Survey,  formerly 
sent  me  specimens  from  the  following  localities :  Sooke,  Vancouver 
Sound,  one  very  large,  21  inches  across  (Macoun,  1887)  ;  Barclay 
Sound,  one  very  large,  20  inches  across  (1887)  ;  Discovery  Sound, 
one  large  six-rayed  example,  otherwise  normal  (G.  M.  Dawson, 
1885)  ;  Queen  Charlotte  Islands  (Dawson,  1885). 

I  have  also  examined  many  specimens  from  Victoria,  B.  C. ;  Puget 
Sound;  Tomales  Bay;  Monterey;  San  Luis  Obispo  Bay;  San 
Diego,  Calif.,  and  various  other  places  on  the  coast  of  California. 
Its  southern  range  extends  certainly  to  Santa  Barbara,  Calif.,  and 
probably  beyond  San  Diego. 

This  can  scarcely  be  confounded  with  any  other  species,  for  its 
conspicuously  acervate  and  reticulate,  capitate  dorsal  spines  are 
characteristic.  The  reticulation  of  E.  troschelii  and  L.  epichlora 
is  much  less  conspicuous,  and  the  former  has  more  slender  rays; 
but  both,  even  when  very  young,  differ  in  having  biserial  adambu- 
lacral  spines.  Larger  specimens  could  hardly  be  mistaken. 


72  .  /  VERRILL 

PISASTER  OCHRACEUS,  Var.  NODIFERUS  Verrill,  nov. 
Plate  LVI,  figures  3,  30. 

This  narne  is  proposed  for  that  variety  of  this  species  which  has 
the  dorsal  spines  collected  into  large  detached  groups. 

The  form  is  the  same  as  in  the  typical  reticulated  variety,  and 
specimens  occur  of  all  sizes  from  two  inches  up  to  a  foot  or  more  in 
diameter. 

A  strongly  characterized  small  specimen  from  Monterey  has  the 
radii  16  mm.  and  52  mm. ;  ratio,  1 : 3.25.  The  dorsal  ossicles  are 
reticulate,  but  so  many  are  without  spines  that  the  latter  show  no 
regular  reticulate  pattern.  The  Geological  Survey  of  Canada  sent  me 
several  specimens  with  the  nodular  character  even  more  prominent. 

The  dorsal  spines  are  all  capitate  and  somewhat  unequal;  they 
form  a  conspicuous  pentagon  with  a  central  cluster  on  the  disk  and 
five  prominent  median  radial  rows  proximally.  Between  the  median 
and  superomarginal  rows  there  are  ten  to  twelve  large  irregularly 
placed  and  very  prominent  clusters  of  capitate  spines,  with  about 
five  to  eight  spines  in  each  group. 

The  superomarginal  spines  form  a  simple  regular  row  in  the  type ; 
they  are  about  equal  in  size  to  the  dorsal  spines,  but  more  conical. 
The  ventral  and  adambulacral  spines  and  the  pedicellariae  are  like 
those  of  the  typical  form.  A  very  young  specimen,  also  from  Mon- 
terey, has  the  radii  8  mm.  and  24  mm. ;  ratio,  i :  3. 

This  small  one  agrees  with  the  preceding  in  all  essential  characters, 
but  the  spines  are  correspondingly  smaller  and  there  are  but  two 
distinct  rows  of  interactinal  spines.  The  pentagon  on  the  disk  is  well 
marked. 

This  variety  occurs  commonly  from  Monterey,  Calif.,  north  to 
Sitka,  Alaska.  It  is  the  predominating  variety  on  some  parts  of  the 
coast  of  British  Columbia,  on  rocks  at  low  tide. 

Six-rayed  specimens  are  not  very  rare. 

HYBRIDS  ? 

There  are  indications  that  this  species  may  hybridize  with 
L.  epichlora  and  with  others,  for  where  they  live  associated,  as  at 
Sitka,  specimens  occur  that  seem  to  belong  to  Leptasterias  epichlora, 
but  have  some  of  the  characters  of  ochraceus,  in  a  greater  or  less 
degree. 

One  young  L.  epichlora  from  Wrangel,  Alaska  (var.  subnodu- 
losa),  has  only  five  rays,  and  the  dorsal  spines  are  more  strongly 
acervate  and  much  larger  than  usual,  giving  it  a  close  superficial 


SHALLOW-WATER   STARFISHES  73 

resemblance  to  the  young  of  ochraceus,  var.  nodiferus,  of  similar  size 
(diameter  about  4  inches),  but  it  has  the  biserial  adambulacral 
spines  and  other  characters  of  epichlora,  and  lacks  the  giant  serrate- 
pedicellariae.     (See  under  L.  epichlora,  var.  subnodulosa.) 

Other  specimens  have  these  variations  in  less  degree,  and  many 
have  more  or  less  of  the  giant  lateral  pedicellariae,  which  may  indi- 
cate an  infusion  of  ochraceus  blood,  so  to  speak,  for  they  are  not 
constantly  present  in  epichlora  and  do  not  exist  in  the  allied  species. 

Where  so  many  closely  related  species  are  associated  together,  as 
on  the  Alaskan  coasts,  occasional  hybridization  is  to  be  expected.  It 
is  well  known  to  occur  on  certain  middle  portions  of  the  New  Eng- 
land coast,  between  A.  vulgaris  and  A.  forbesi,  where  the  ranges  of 
these  two  species  overlap,  as  well  as  their  breeding  seasons,  as  at 
Wood's  Hole  and  Vineyard  Sound,  Mass.,  just  south  of  Cape  Cod. 

The  Asterias  janthina  Brandt  (Prod.,  p.  269,  1835),  probably 
described  from  a  colored  drawing  only,  was  so  poorly  characterized 
that  no  one  has  been  able  to  identify  it  with  certainty.  Dr.  Stimpson 
thought  that  it  might  be  only  a  color-variety  of  ochraceus,  and  that 
is  not  unlikely,  from  Brandt's  statement.  His  description  was  as 
follows : 

"A  species  quite  similar  to  the  preceding  (ochraceus)  but  dif- 
ferent. Diameter  of  the  disk  less,  rays  a  little  longer,  with  more 
numerous  and  more  crowded  spines,  much  more  numerous  in  the 
center  of  the  disk,  and  not  forming  a  pentagonal  star.  Back  all  light 
violaceus ;  spines  white." 

The  relative  length  of  the  rays  in  P.  ochraceus  is  quite  variable, 
and  so  is  the  number  and  arrangement  of  the  dorsal  spines.  Some 
of  our  Sitka  and  Vancouver  Island  specimens  have  the  spines  closely 
crowded.  The  color  in  most  species  of  this  group  may  vary  between 
yellow,  orange,  red,  and  purple.  This  species  is  often  violet  or 
purple,  in  life. 

PISASTER  CONFERTUS  (Stimpson). 
Plate  xxxvin,  figures  i,  2  (type)  ;  plate  LIII,  figure  2. 
Asterias  conferta  STIMPSON,  Proc.  Boston  Soc.  Nat.  History,  vm,  p.  263,  1862. 
Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  p.  326,  1867.     Perrier,  Revision,  p. 
335  [7i]»  1875.     Sladen,  op.  clt.,  p.  820.    Bell.  Proc.  Zool.  Soc.  London, 
p.  494,  1881.    Whiteaves,  Trans.  Royal  Soc.  Canada,  iv,  p.  116,  1887.    De 
Loriol,  Mem.  Soc.  Phys.  et  Hist.  Nat.,  Geneve,  xxxii,  part  2,  p.  17,  pi. 
m,  figs,  i-ig,  1897. 

Dr.  Stimpson's  original  description  was  as  follows: 

"  Rays  five,  stout,  rounded,  and  dilated  at  base ;  disc  large.    Pro- 


74  VERRILL 

portion  of  smaller  to  greater  diameter,  i :  3.5.  Ambulacral  furrows 
broad  at  the  base  of  the  ray,  where  the  pores  are  crowded  into  six 
rows.  Ambulacral  spines  in  one  regular  row,  slender,  compressed ; 
subcylindric,  as  long  as  the  ventral  spines,  somewhat  tapering,  with 
bluntly-rounded  tips.  Ventral  spines  crowded,  very  numerous,  in 
six  or  seven  rows  near  the  base  of  the  ray ;  short,  subcylindric,  a  little 
bent  outward,  and  slightly  flattened  externally,  with  more  or  less 
longitudinal  striation  near  the  tip.  Beyond  the  ventral  spines  there 
are  no  regular  channels,  but  the  surface  is  covered  with  pedicellariae 
and  minute  spines.  The  dorsal  spines  are  numerous,  equal  in  size, 
but  very  small, — not  more  than  one-twentieth  of  an  inch  in  length, 
scarcely  capitate,  but  more  or  less  pointed.  They  are  uniformly  dis- 
tributed over  the  disc  and  rays  in  a  reticulating  manner,  the  inter- 
spaces being  from  one-tenth  to  one-half  inch  in  diameter,  and 
thickly  crowded  with  groups  of  papulae  and  minor  pedicellariae ;  the 
latter  generally  occurring  about  the  bases  of  the  spines,  but  not 
forming  wreaths  or  crowns.  The  major  pedicellariae  are  very  short 
and  stout,  regularly  conical  or  somewhat  wedge-shaped,  with  very 
broad  valves.  They  are  very  numerous,  particularly  on  the  disc, 
where  they  often  stand  in  groups  of  ten  to  twenty  together.  The 
largest  are  found  isolated  in  the  angles  of  the  rays  below,  where  one 
occurred  having  a  diameter  of  eight-hundredths  of  an  inch,  and  a 
regularly  conical  form,  with  a  square  base.  Diameter  of  our  only 
specimen,  ten  inches. 

"  It  is  allied  to  A.  ochracea,  but  differs  in  its  more  numerous 
spines,  pedicellariae,  and  ambulacral  pores. 

"  Habitat,  Puget  Sound — North  West  Boundary  Commission.  Dr. 
C.  B.  Kennerly." 

I  have  reproduced  large  photographs  of  the  type  of  this  species, 
now  in  the  National  Museum.  These  were  sent  by  Dr.  R.  Rathbun. 
(See  pi.  xxxvin,  figs,  i,  2,  reduced.) 

In  this  specimen  the  radii  are  about  45  mm.  and  162  mm. ;  ratio, 
about  as  i :  3.6.  The  rays  are  thick  at  base  and  taper  rapidly,  though 
they  differ  considerably  in  this  respect,  indicating  that  the  skeleton 
in  life  is  rather  flexible  and  weak.  The  crowded  six-rowed  or  eight- 
rowed  arrangement  of  the  proximal  ambulacral  pores  is  very  evident. 
The  slender  adambulacral  spines  near  the  mouth  are  much  longer 
than  those  farther  out,  as  in  some  other  species;  they  stand  singly 
on  the  plates,  in  one  regular  row.  The  ventral  spines  are  pretty  uni- 
form in  size  and  form,  and  are  nearly  smooth  and  blunt ;  they  stand 
mostly  in  six  rows  proximally,  but  form  only  four  or  five  rows  at  the 


SHALLOW-WATER   STARFISHES  75 

middle  of  the  ray.    Probably  there  are  at  least  four  rows  of  inter- 
actinal  ossicles. 

The  dorsal  spines,  as  stated,  are  small,  mostly  subacute,  and 
arranged  in  a  distinctly  areolated  or  reticulated  pattern,  many  of 
them  forming  regular  transverse  or  oblique  rows  or  combs  on  the 
reticulating  ossicles;  but  they  differ  rather  more  in  size  than  indi- 
cated in  the  original  description,  some  of  them  being  much 
smaller,  more  slender  and  very  acute.  But  they  are  far  more  uni- 
form than  in  P.  ochraceus.  They  do  not  form  a  definite  median  row, 
The  dermal  minor  pedicellariae  are  numerous.  The  dorsal,  sessile, 
dermal  major  pedicellariae  are  often  larger  than  the  spines  on  the 
dorsal  surface,  especially  at  the  interbrachial  angles  above,  and  also 
below,  and  along  the  lateral  channels.  The  actinal  spines  bear  thick 
clusters  of  minor  pedicellariae  on  their  outer  sides. 

VARIATIONS. 

A  specimen  of  this  species  from  Malaspina  Inlet,  coll.  G.  M. 
Dawson,  1885,  received  from  the  Geological  Survey  of  Canada 
through  J.  F.  Whiteaves,  agrees  well  with  the  type  in  most  respects, 
but  it  has  distinct  radial  bands  of  small  spines.  Radii,  28  mm.  and 
no  mm.;  ratio,  1:4;  but  the  dried  specimen  is  considerably  flat- 
tened, due  to  imperfect  preservation. 

The  dorsal  surface  is  very  irregularly  reticulated;  the  numerous 
small  spines  have  a  tendency  in  many  parts  to  form  rather  long 
transverse  or  oblique  rows.  Most  of  the  spines  are  short  and  blunt, 
or  subacute,  but  some  of  the  smaller  ones  are  slender  and  very  acute. 
Minor  pedicellariae  of  very  small  sizes  are  exceedingly  abundant, 
both  in  wreaths  around  the  bases  of  the  spines  and  on  all  the  inter- 
mediate surface.  The  superomarginal  spines  form  a  somewhat 
irregular  band  of  small  spines,  scarcely  larger  than  the  dorsals,  sev- 
eral standing  on  each  ossicle.  The  inf  eromarginal  spines  are  similar, 
a  trifle  longer,  blunt,  and  mostly  striated  distally,  two  often  standing 
on  one  plate.  The  interactinal  spines,  which  form  three  or  four 
rows  proximally,  are  similar  in  form,  but  increase  in  size  toward  the 
grooves.  The  adambulacral  spines  are  long,  slender,  and  regularly 
tapered.  The  giant  pedicellariae  occur  mostly  on  the  interradial 
areas  and  lateral  channels.  Some  of  them  are  stouter  than  the  inter- 
actinal spines,  but  shorter.  They  are  usually  stout  and  blunt,  very 
obtusely  conical  or  blunt  wedge-shaped,  with  slightly  denticulate 
tips.  The  ambulacral  pores  are  crowded  into  eight  rows  proximally. 


76  VERRILL 

M.  de  Loriol  (op.  cit,  1896,  p.  17,  pi.  in,  figs,  i-i^)  has  described 
and  figured  a  specimen,  evidently  of  this  species,  from  Vancouver 
Island.  But  his  general  figure  (i)  makes  the  dorsal  reticulation 
much  more  regular  than  in  the  type,  and  the  madreporic  plate  is  put 
in  the  center  of  the  disk,  which  is  probably  the  fault  of  the  artist. 

I  have  received  several  other  very  similar  specimens  from  near 
Victoria  (C.  F.  Newcombe,  coll.)  and  from  Departure  Bay  (Geo- 
logical Survey  of  Canada).  None  of  these  agree  perfectly  with  the 
type. 

This  species  is  rare  in  collections  and  seems  to  be  known  only  from 
Puget  Sound,  Vancouver  Island,  and  British  Columbia,  where  it  is 
said  to  be  common. 

It  is  closely  related  to  P.  ochraceus  and  P.  fissipinus  by  the  char- 
acter of  the  ventral  and  adambulacral  spines,  and  by  the  reticulated 
dorsal  surface.  But  the  dorsal  spinulation  is  even  more  like  that  of 
E.  acanthostoma  V.  The  latter  has  much  longer  rays,  a  much 
feebler  skeleton,  two  rows  of  adambulacral  spines,  and  lacks  the  very 
large,  dentate,  dermal  pedicellariae. 

PISASTER  FISSISPINUS  (Stimpson). 
Plate  xxxix,  figures  I,  2  (type). 

Asterias  fissispina  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vin,  p.  264,  1862. 

Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  p.  326,  1867. 
Pisaster  fissispina  VERRILL,  Amer.  Journ.  Sci.,  xxvm,  p.  63,  1909. 

Dr.  Stimpson's  original  description  is  as  follows: 
"  Five  rays,  short,  and  dilated  at  base ;  disc  large.  Proportion  of 
the  diameters,  i :  3.  Ambulacral  pores  near  base  of  ray  crowded, 
alternating,  indistinctly  six  or  eight  rowed.  Ambulacral  spines  in 
one  regular  row,  as  long  as  the  ventrals,  and  flattened  on  the  outer 
side.  Ventral  spines  sub-equal,  stout,  sub-cylindrical,  truncated,  with 
fissured  tips,  and  a  deep,  longitudinal  sulcus  on  the  outer  side ;  they 
form  five  regular  rows.  The  marginal  dorsal  spines  are  as  large  as 
the  ventrals,  capitate,  with  striated  sides  and  pinched  tips,  and  form 
an  irregular  row,  of  much  fewer  spines  than  occur  in  a  ventral  row. 
The  spines  of  the  back  are  few  in  number,  and  of  only  half  the  size 
of  the  ventral  spines.  They  are  shaped  like  the  marginal  spines, 
from  which  they  are  not  ordinarily  distinct,  and  are  arranged  on 
reticulating  ridges,  forming  a  rather  open  net-work.  On  the  disc 
they  form  a  pentagon,  from  each  angle  of  which  extends  a  median 
row  reaching  to  one-third  the  length  of  the  ray.  Within  the  penta- 
gon, close  to  its  periphery,  the  madreporic  plate  is  situated.  The 


SHALLOW-WATER   STARFISHES  77 

dorsal  spines  increase  in  size  toward  the  tips  of  the  rays.    Papulae 
ing  groups.    Minor  pedicellariae  nearly  as  in  A.  ochracea  and  A.  con- 
fer ta;  major  pedicellariae  smaller  and  far  less  numerous  than  in    - 
those  species,  but  of  similar  short  and  stout  form.    Diameter,  thir- 
teen inches. 

"  Habitat  Shoalwater  Bay,  Oregon  Coast.  Northern  Pacific  Rail- 
road Expedition.  Dr.  J.  G.  Cooper." 

Large  photographs  of  the  type  of  this  species,  which  is  still  in  the 
U.  S.  National  Museum,  were  furnished  by  Dr.  R.  Rathbun.  They 
indicate  that  the  average  radii  are  60  mm.  and  170  mm.;  ratio, 
about  as  1 : 2.83.  The  rays  taper  rapidly  from  a  broad  disk. 

The  slender  adambulacral  spines  are  compressed  and  much 
crowded,  one  to  a  plate.  The  peractinal  and  synactinal  spines  are 
mostly  gouge-shaped,  though  often  bifid  at  tip ;  but  on  the  distal  half 
of  the  ray,  many  are  only  sulcate  and  flattened.  The  dorsal  surface 
was  very  accurately  described  by  Dr.  Stimpson,  as  may  be  seen  by 
comparison  with  our  figures  of  his  type. 

This  species  is  more  nearly  allied  to  P.  ochraceus  than  to  any 
other.  It  differs  in  the  peculiar  gouge-shaped  spines,  in  the  more 
open  reticulation  of  the  dorsal  skeleton,  and  in  the  much  smaller  and 
more  slender  dorsal  spines. 

The  locality  given  by  Stimpson  is  the  only  one  positively  known 
to  me  for  this  species. 

PISASTER  BREVISPINUS  (Stimpson). 

Plate  XLI,  figures  i,  2  (type)  ;  plate  XLIV,  figures  I,  2 ;  plate  XLV,  figure  i ; 

plate  LXIX,  figure  3;  plate  LXXVI,  figures  i-ib  (details). 
Asterias  brevispina  STIMPSON,  Journ.  Boston  Soc.  Nat.  Hist,  vi,  p.  88,  pi. 

xxm,  fig.  3  (ventral  spines),  1857. 
Pisaster  brevispina  VERRILL,  Amer.  Journ.  Sci.,  xxvm,  p.  63,  1909. 

The  original  description  by  Dr.  Stimpson  is  as  follows : 
"  Rays  five,  each  equalling  in  length  twice  the  diameter  of  the  disk. 
Upper  surface  covered  with  very  short,  blunt,  nearly  uniform  spines, 
moderately  numerous,  sometimes  forming  an  irregular  row  along 
the  middle  of  the  ray,  and  showing  a  tendency  to  reticulation  on  the 
sides.  Beneath,  there  is  a  single  row  of  slender  ambulacral  spines, 
which  are  blunt  and  somewhat  irregular  in  length;  between  these 
and  the  marginal  channel  there  are  four  rows  of  short  compressed 
spines,  gouge-shaped,  or  notched  by  an  oblique  concavity  at  their 
truncated  extremities.  Madreporic  body  large.  Color  yellowish. 
Diameter,  six  inches.  Taken  from  a  sandy  bottom  in  ten  fathoms 
near  the  mouth  of  San  Francisco  Bay." 


78  VERRILL 

Dr.  Rathbun  has  sent  nearly  natural-size  photographs  of  the 
original  type  of  this  species,  which  seems  to  be  almost  unknown  in 
many  later  collections. 

The  disk  is  of  rather  large  size  and  elevated ;  the  five  rays  taper 
rapidly  from  rather  wide  and  swollen  bases.  The  radii  are  about 
30  mm.  and  116  mm.,  if  the  photographs  are  correctly  marked  as  to 
reduction ;  ratio,  about  as  i :  3.87. 

The  adambulacral  spines  are  small  and  slender,  mostly  compressed, 
crowded  in  one  row,  one  to  a  plate ;  those  near  the  mouth  are  longer. 

The  inf  eromarginal  and  actinal  spines  are  very  much  larger,  rather 
short  and  very  stout,  mostly  clavate  or  subspatulate,  and  with  a 
deeply  grooved  or  gouge-shaped  tip ;  but  many  are  simply  sulcate  and 
little  flattened.  They  form  four  or  five  crowded  rows  near  the  bases 
of  the  rays,  the  inferomarginals  bearing  two  spines,  but  at  about 
the  middle  they  are  reduced  to  three  or  four  regular  rows.  Most  of 
them  bear  close  clusters  of  minor  pedicellariae  on  their  outer  side. 
The  superomarginal  spines  are  smaller,  fusiform  or  subconical,  sub- 
acute;  they  form  a  pretty  regular  row,  separated  from  the  infero- 
marginals by  a  rather  wide  channel. 

The  dorsal  spines  are  short,  thick,  nearly  equal,  with  conical, 
mostly  subacute  tips.  They  are  rather  irregularly  distributed,  in 
short  broken  rows  or  singly ;  they  form  a  somewhat  evident  median 
radial  series,  of  two  or  three  close  irregularly  alternating  rows  on 
the  basal  part  of  the  rays ;  on  the  sides  many  of  them  are  in  obliquely 
transverse  short  rows,  and  sometimes  short  longitudinal  rows  can 
be  traced.  They  bear  small  basal  groups  of  minor  pedicellariae. 
Dermal  minor  pedicellariae  everywhere  scattered  between  the  spines. 
The  sessile,  dermal,  major  pedicellariae  appear  to  be  few  and  mostly 
along  the  submarginal  channel;  they  are  rather  small,  short,  sub- 
conic,  obtuse.  A  few  lanceolate,  acute,  major  pedicellariae  occur 
along  the  edges  of  the  ambulacral  grooves. 

VARIATIONS. 

This  species  varies  considerably  in  appearance,  according  to  age, 
locality,  and  mode  of  preservation.  When  dried  carefully,  from 
alcohol,  the  disk  is  high  and  convex,  while  the  mouth  and  jaws  are 
deeply  sunken;  the  rays  are  stout  and  high  at  base  but  rapidly 
tapered.  There  is  usually  a  distinct  median  dorsal  row  of  spines, 
simple  in  the  younger  specimens,  but  often  double-  or  triple-ranked 
in  the  older  ones. 


SHALLOW-WATER   STARFISHES  79 

On  the  medium-sized  specimens  the  pentagon  of  the  disk  is  very 
evident,  but  in  large  specimens  it  is  obscured  by  the  numerous  inter- 
polated scattered  spines.  In  those  about  six  to  eight  inches  in 
diameter,  the  central  plate,  five  radial,  and  five  interradial  primary 
plates  are  evident.  Each  bears  a  cluster,  usually  of  four  to  six  short, 
thick,  clavate  spines,  with  ovate  or  acorn-shaped  striated  tips.  They 
are  similar,  in  size  and  form,  to  those  of  the  median  radial  rows. 

Other  somewhat  smaller,  but  similar,  spines  are  scattered  within 
and  around  the  primary  pentagon,  and  over  the  dorsal  surface  gen- 
erally. But  the  dorsal  spines  are  not  numerous  in  the  young.  They 
do  not  form  distinct  rows,  but  are  irregularly  scattered,  and  are 
mostly  pretty  nearly  equal  in  size  and  form.  In  the  larger  specimen, 
ten  to  twelve  inches  in  diameter,  the  dorsal  spines  become  decidedly 
more  numerous,  and  many  are  grouped  two  to  four  or  more  on  one 
plate. 

The  superomarginal  spines  are  similar  to  the  dorsals,  usually  with 
rather  more  conical  tips,  which  are  strongly  striated.  They  form  a 
regular,  close  row,  simple  in  the  smaller  specimens,  but  often  double 
proximally  in  the  larger  ones. 

The  inferomarginal  row  is  very  regular  and  nearly  always  double, 
two  spines  standing  obliquely  on  each  plate,  sometimes  three  in  the 
larger  specimens.  They  are  rather  stouter  and  more  obtuse  than  the 
upper  marginals,  usually  with  the  tips  striated  in  small  specimens; 
but  in  the  larger  ones  these  spines  mostly  lose  their  striations,  and 
many  become  more  or  less  flattened  distally,  while  some  are  apt  to 
have  a  distal  furrow  on  the  upper  side  ("gouge-shaped"),  though 
this  is  by  no  means  constant. 

In  nearly  all  the  specimens  there  are  but  two  simple,  regular 
actinal  rows  of  spines,  making,  with  the  inferomarginals,  four  close, 
unusually  regular  rows.  The  actinals  are  short  and  stout,  similar 
to  the  inferomarginals.  In  the  larger  specimens  some  of  them  are 
somewhat  flattened,  and  some  are  often  grooved  or  slightly  bilobed 
distally. 

The  adambulacral  spines  are  slender,  four  or  sometimes  five  corre- 
sponding to  each  synactinal  plate.  In  young  examples  they  are 
mostly  regularly  tapered  and  form  a  very  regular  even  row ;  in  the 
larger  ones,  many  of  them  are  often  stouter,  obtuse,  cylindric,  a 
little  flattened,  or  even  slightly  clavate.  They  become  longer  proxi- 
mally, near  the  mouth. 

Major  pedicellariae  of  rather  large  size  occur  on  the  back,  sides, 
and  interradial  axils,  on  the  adambulacral  spines,  and  on  the  inner 


8O  VERRILL 

margins  of  the  grooves.  Those  of  the  actinal  side  and  ambulacral 
furrows  are  mostly  acute-lanceolate  or  acuminate,  compressed,  with 
the  sharp  curved  tips  often  overlapping.  They  often  exceed  the 
adambulacral  spines  in  diameter.  Those  of  the  dorsal  surface  are 
shorter,  stouter,  ovate,  with  narrow  obtuse  tips,  which  are  slightly 
denticulate.  Those  on  the  intermarginal  furrow  are  largest  and 
stoutest,  with  stronger  denticulations. 

Minor  pedicellariae  are  very  small.  They  form  small  wreaths 
around  most  of  the  dorsal  spines,  and  clusters  on  the  outside  of  the 
ventral  ones;  small  clusters  also  occur  on  the  inside  margin  of  the 
adambulacral  plates  and  on  some  the  larger  major  pedicellariae.  On 
the  dorsal  surface  they  form  mostly  rather  small  clusters,  pretty 
evenly  scattered  over  the  whole  surface;  in  large  specimens  they 
often  become  very  numerous  and  pretty  evenly  distributed. 

The  papulae  are  very  small  and  apparently  not  very  numerous, 
except  in  large  examples.  The  integument  is  thick  and  firm,  so  that 
it  conceals  the  outlines  of  the  plates. 

The  mouth  is  deeply  sunken.  The  jaws  are  elongated,  narrow  or 
much  compressed,  nearly  perpendicular  within;  the  adoral  spines 
form  two  close  rows  of  about  eight  each ;  the  apical  oral  spines  are 
rather  stouter  than  the  rest ;  there  is  often  but  one  to  a  jaw. 

The  color,  in  the  only  case  noted,  was  pale  purple  or  pink. 

A  good  specimen  from  off  Monterey,  Calif.  (No.  1829^  M.  C.  Z.), 
differs  somewhat  from  all  the  others.  Radii,  33  mm.  and  130  mm. ; 
height  in  middle,  40  mm. 

The  central  and  ten  primary  radial  and  interradial  plates  are 
covered  with  clusters  of  short  capitate  spines  with  obtuse  tips,  about 
eight  to  ten  in  a  cluster.  About  ten  smaller  plates  within  the  pen- 
tagon are  indicated  by  small  clusters  of  simple  spines.  The  median 
radial  rows  are  conspicuous.  On  the  proximal  third  of  the  rays  they 
mostly  contain  three  regular  rows  of  spines;  further  out  the  three 
rows  are  less  regular.  On  the  distal  fourth  there  is  usually  but  one 
row.  The  other  dorsal  spines  are  mostly  grouped  in  clusters  of  three 
to  six,  the  clusters  being  irregularly  scattered,  largest  on  the  sides 
of  the  rays.  The  spines  are  all  similar,  short,  thick,  with  more  or 
less  conical,  obtuse  or  ovate  striated  tips.  The  marginal  and  actinal 
rows  of  spines  are  very  regular  and  even.  They  stand  in  close  rows. 
There  is  a  wide  and  deep  intermarginal  channel.  The  superomar- 
ginal  row  of  spines  is  simple.  The  inferomarginal  row  is  double. 
The  two  interactinal  rows  are  simple  and  nearly  equal. 


SHALLOW-WATER   STARFISHES  8l 

MEASUREMENTS. 

Radii 
Serial  number.         'Lesser.  Greater.  Locality. 

I304a  SO  mm.  168  mm.  Off  San  Francisco.                %dUv£<^w^v«    k*t  j 

13046  26  mm.  73  mm.  Off  San  Francisco. 

13040  18  mm.  60  mm.  Off  San  Francisco. 

1304^  42  mm.  no  mm.  Off  San  Francisco. 

18216  22  mm.  63  mm.  Off  Santa  Cruz,  Calif. 

18200  33  mm.  130  mm.  Off  Monterey,  Calif. 

This  species  ranges  from  south  of  San  Francisco  to  Puget  Sound 
and  northward  to  Departure  Bay,  B.  C.  In  the  Museum  of  Comp.  V*' 
Zoology  at  Cambridge,  Mass.,  I  have  studied  a  good  series  from  off 
San  Francisco  (No.  1304)  ;  Crescent  City,  Calif.  (No.  1303^)  ;  Gulf 
of  Georgia,  A.  Agassiz  (No.  1301)';  Monterey  Bay  (No.  1820,  see 
figs.);  off  Santa  Cruz  I.,  Calif.  (No.  1821).  It  appears  to  be 
common  in  shallow  water  on  the  Californian  coast.  Very  few 
authentic  localities  have  been  recorded  hitherto,  except  that  given  by 
Dr.  Stimpson  (Bay  of  San  Francisco).  A  single  large  specimen 
has  been  sent  to  me  from  Departure  Bay,  B.  C.  (coll.  H.  C.  Young,  *CjU^fa*rH.m  (_ 
Geological  Survey  of  Canada). 

PISASTER  CAPITATUS  (Stimpson). 
Plate  xxxvi,  figures  3,  4  (type)  ;  plate  LVI,  figure  4. 

Asterias  capitata  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vm,  p.  264,  1862. 
Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  p.  327,  1867.  ?  Perrier,  Arch.  Zool. 
Exper.,  rv,  p.  335,  1875  (six-rayed  specimen,  no  descr.). 

Dr.  Stimpson  described  this  species  as  follows: 
"  Rays  five,  not  contracted  at  base ;  disc  large.  Proportion  of 
diameters,  i :  4.5.  Ambulacral  pores  rather  narrow,  in  four  regular 
rows.  Ambulacral  [adambulacral]  spines  in  one  regular  row,  linear, 
compressed,  and  blunt.  Ventral  spines  as  long  as  the  ambulacrals, 
capitate,  with  bluntly-rounded  heads,  elegantly  striated  on  the  con- 
vex inner  face  and  tip,  and  with  a  median  sulcus  on  the  outer  side. 
They  are  arranged  in  four  rows,  those  of  the  outer  row  being  largest ; 
and  there  are  some  minor  pedicellariae  on  the  outer  sides  of  the 
spines  in  all  of  the  rows.  The  dorsal  spines  are  not  very  numerous, 
but  are  for  the  most  part  large,  their  regularly  globular  and  beauti- 
fully striated  or  radiated  heads  being  about  eight-hundredths  of  an 
inch  in  diameter,  and  larger  than  those  of  the  ventral  spines.  They 
are  arranged  without  order,  standing  about  one-seventh  of  an  inch 
apart;  but  five  or  six  longitudinal  rows  may  be  obscurely  traced, 
the  marginal  row  being  most  distinct,  containing  eighteen  or  twenty 

7 


82 


VERRILL 


spines.  On  the  disc,  there  is  a  central  tubercle,  but  scarce  any  indi- 
cations of  a  pentagon.  Around  the  bases  of  the  dorsal  spines  there 
are  regular  wreaths  of  minor  pedicellariae.  The  major  pedicellariae, 
which  are  of  the  short,  conical  or  sub-globular  form,  with  broad 
valves,  are  scattered  between  the  spines  as  in  A.  ochracea.  Papulae 
in  groups. 

"  Color  in  life  purple,  according  to  Dr.  Newberry.  The  spines 
are  probably  white,  or  at  least  of  a  lighter  color.  Diameter,  five  and 
a  half  inches. 

"This  very  pretty  species  differs  from  A.  ochracea  in  its  larger 
dorsal  spines,  which  are  not  arranged  in  a  reticulating  pattern ;  and 
from  A.  Lutkenii  in  its  shorter  and  more  numerous  ventral  spines, 
as  well  as  in  the  presence  of  major  pedicellariae  on  the  back. 

"  Habitat,  San  Diego,  Cal.  Colorado  Expedition.  Dr.  J.  S.  New- 
berry." 

The  photographs  of  the  type,  sent  by  Dr.  R.  Rathbun,  are  six 
inches  in  longest  diameter ;  radii,  18  mm.  and  80  mm. ;  ratio,  nearly 
1:4.5. 

Little  need  be  added  to  Stimpson's  description  of  the  type.  The 
adambulacral  spines  are  often  flattened.  The  ventral  spines  are 
unequal ;  those  of  the  inner  or  synactinal  row  are  distinctly  smaller 
than  the  others,  and  the  outer  or  peractinal  are  largest.  Most  of 
these  spines  are  stout  and  clavate,  rather  than  capitate,  the  tips  being 
mostly  obtusely  conical  or  ovoid  and  strongly  striated;  only  a  part 
are  grooved ;  distally  they  become  much  more  slender  and  distinctly 
clavate. 

The  superomarginal  spines  are  considerably  stouter  than  the 
ventral  ones,  but  not  much  longer.  They  form  a  regular  simple  row 
of  rather  widely  spaced  spines,  each  one  on  the  raised  node  of  a 
marginal  ossicle.  The  dorsal  spines  are  mostly  quite  similar  to  the 
laterals  in  size  and  form.  The  larger  ones  have  large,  rounded, 
capitate,  and  strongly  striate  tops ;  but  those  toward  the  ends  of  the 
rays  become  longer  and  more  slender,  cylindrical  or  clavate,  rather 
than  capitate,  but  striated  in  the  same  way. 

The  spines  are  distantly  spaced,  generally  one  on  each  raised 
node  of  the  reticulating  ossicles.  Usually,  in  our  specimens,  five  or 
six  ridges,  indicating  the  lobes  of  the  ossicles,  radiate  out  from  the 
base  of  each  spine,  so  that  the  skeleton  has  a  pretty  regularly  reticu- 
lated structure.  The  spines  usually  form  more  or  less  evident 
median,  radial  rows,  and  sometimes  two  less  distinct  rows  can  be 
traced,  between  the  median  and  marginal  rows. 


SHALLOW-WATER  STARFISHES  83 

In  the  Museum  of  Yale  University  there  is  a  specimen  of  this 
species  collected  at  Monterey,  Calif.,  which  agrees  very  closely  with 
the  type  in  size  and  structure. 

The  radii  are  20  mm.  and  70  mm. ;  ratio,  1 : 3.5.  The  rays  are 
shorter  than  in  the  type.  The  spinulation  agrees  in  all  essential 
points  with  the  type,  though  the  distal  dorsal  spines  of  the  rays 
appear  longer  and  less  stout,  and  more  of  them  have  the  clavate  form. 

The  wreaths  of  minor  pedicellariae  about  the  spines  are  not  con- 
spicuous, but  they  contain  very  large  numbers  of  very  minute  pedi- 
cellariae. Others  of  similar  small  size  are  scattered  between  the 
spines  or  grouped  in  dermal  clusters  on  or  around  the  large  papular 
areas.  The  papulae  are  small  and  form  large  groups,  above  and 
below. 

The  dorsal  and  lateral  dermal  major  pedicellariae  are  moderately 
large  and  stout,  ovoid  or  subconic,  with  obtuse  and  usually  denticu- 
lated jaws.  They  are  not  nearly  so  numerous  nor  so  large  as  in  the 
type.  Most  of  the  actinal  spines  are  clavate  and  many  lack  the  sulcus 
on  the  outside;  the  synactinal  spines  are  smaller  and  more  cylin- 
drical; between  them  are  large  papular  areas.  The  dorsal  skeleton 
is  firm  and  distinctly  reticulated,  with  a  tendency  to  a  six-rayed 
arrangement  around  the  principal  spines,  which  are  not  so  numerous 
as  in  the  type. 

Monterey,  Calif.  (R.  E.  C.  Stearns,  Mus.  Yale  Univ.) ;  San 
Diego  (Stimpson,  Ives,  and  others)  ;  San  Diego  (Mus.  Yale  Univ. 
coll.). 

This  species  is  closely  allied  to  P.  liitkenii.  It  differs  especially 
in  having  much  fewer  dorsal  spines,  which  are  also  larger  and  more 
distinctly  capitate.  The  ventral  (actinal)  spines  are  more  unequal, 
stouter,  shorter,  and  more  capitate. 

PISASTER  LUTKENII   (Stimpson). 

Jl*- 
Plate  XL,  figures  I,  2  (type). 

Asterias  liitkenii  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vm,  p.  265,  1862. 
Verrill,  op.  cit.,  1867,  p.  32.  Perrier,  Revis.,  p.  70,  1875.  Bell,  Genus 
Asterias,  Proc.  Zool.  Soc.  London,  1881,  p.  495.  Sladen,  op.  cit.,  pp.  566, 
824,  1889.  De  Loriol,  Mem.  Soc.  Phys.  et  Hist.  Nat,  Geneve,  xxxn,  p. 
15,  pi.  ii  [xvn],  figs,  i-i/t,  1897  (description). 
Pisaster  liitkenii  VERRILL,  op.  cit.,  p.  63,  1909.  1*9$ 

Asterias  exquiseta  DE  LORIOL,  Rev.  Zool.  Suisse,  1888,  p.  403,  pi.  xvin.  fig.  2 
(young). 

When  full-grown  this  is  one  of  the  largest  starfishes  on  the  coast. 
Dr.  Stimpson's  type  was  not  half-grown.  The  following  is  his 
description : 


84  VERRILL 

"  Rays  five,  rather  broad,  with  blunt  tips ;  disc  moderately  large. 
Proportion  of  the  diameters,  1 : 4.25.  Ambulacral  pores  in  four 
regular  rows.  Ambulacral  [adambulacral]  spines  in  one  regular 
row,  slender,  long  (but  not  longer  than  the  ventrals),  not  com- 
pressed, but  tapering  to  a  blunt  point.  Ventral  spines  in  four 
approximated  rows,  elongated,  scarcely  capitate;  heads  elongated, 
subtruncate,  striated  within,  and  often  sulcated  along  the  middle  on 
the  outer  side,  where  there  are  always  semicircular  clusters  of  minor 
pedicellariae.  Beyond  the  ventral  spines  there  are  distinct  lateral 
channels.  The  dorsal  spines  are  uniform  in  size  and  distance,  being 
about  one-fourth  of  an  inch  apart  in  one  specimen,  and  arranged 
without  order,  except  in  the  regular  marginal  row,  which  consists  of 
about  thirty  spines ;  and  they  form  no  pentagon  on  the  disc.  They 
are  one-eighth  inch  high,  and  half  that  in  breadth,  capitate,  with  the 
heads  striated  and  conical,  with  pinched  tips.  Around  the  base  of 
each  there  is  a  rather  narrow  ring  of  minor  pedicellariae.  There  are 
no  major  pedicellarige  scattered  among  the  dorsal  spines,  but  a  few 
occur  in  the  lateral  channels,  similar  to  those  of  A.  ochracea  and 
capitata,  but  of  smaller  size  and  with  narrower  valves.  Papulae  in 
groups.  Diameter,  one  foot.  It  differs  from  A.  gigantea  in  having 
only  five  rays,  and  in  other  particulars. 

"  Habitat,  Coast  of  Oregon.     Mus.  Smithsonian." 

Photographs  of  the  types  of  this  species  were  also  kindly  sent  by 
Dr.  R.  Rathbun  from  the  U.  S.  National  Museum.  As  marked,  the 
photographs  give  the  radii  as  24  mm.  and  96  mm. ;  ratio,  i :  4. 

The  description  by  Dr.  Stimpson  is  so  good,  in  this  case,  that  little 
need  be  added  to  it.  The  upper  marginal  spines  form  a  pretty  regu- 
lar row;  they  are  about  as  large  as  the  ventrals,  but  more  conical, 
and  like  the  dorsal  spines  in  form,  though  rather  larger.  The  median 
dorsal  row  of  spines  is  not  often  very  evident ;  in  some  places  indi- 
cations of  four  or  five  other  longitudinal  rows  can  be  seen  on  either 
side,  so  that  there  may  be  ten  to  twelve  spines  in  a  very  irregular 
transverse  series,  but  there  are  no  distinct  transverse  rows.  On  the 
distal  part  of  the  rays  the  wreaths  of  minor  pedicellariae,  around  the 
spines,  become  larger  and  crowded ;  there  are  also  large  clusters  of 
dermal  minor  pedicellariae  scattered  over  the  whole  dorsal  surface, 
many  of  them  on  the  papular  areas,  besides  many  single  pedicellariae. 

Two  very  large  and  perfect  specimens  of  this  species  are  in  the 
Museum  of  Yale  University,  collected  at  Pacific  Grove,  California, 
by  Professor  W.  R.  Coe,  1901. 


SHALLOW-WATER   STARFISHES  85 

One  of  these  (o)  agrees  very  closely  with  the  type,  except  in 
features  due  to  its  much  greater  size.  Its  radii  are  60  mm.  and 
270  mm. ;  ratio,  i :  4.5. 

The  dorsal  surface  is  nearly  uniformly  covered  with  conical 
spines,  not  forming  definite  rows.  About  twenty-one  to  twenty- 
five  may  be  counted  in  an  irregular  transverse  series  at  the  base  of 
the  rays. 

The  upper  marginal  plates  mostly  bear  two  spines  at  the  base  of 
the  rays,  but  only  one  distally.  The  lower  marginals  bear  two  com- 
plete rows,  to  the  tips  of  the  rays. 

There  are  three  regular  rows  of  actinal  spines  proximally,  mostly 
one  to  a  plate ;  the  first  is  complete  to  the  tips ;  the  second  ceases  near 
the  tip;  the  third  at  about  the  distal  third. 

The  inferomarginal  row  and  first  two  actinals  are  all  close 
together  and  much  alike ;  they  are  rather  short,  very  stout,  clavate, 
with  rounded,  striate  tips,  channeled  on  the  outer  side.  The  syn- 
actinal  spines  are  longer  and  not  quite  so  stout,  tapered  and  chan- 
neled on  the  outer  side.  On  the  interradial  areas  some  of  the 
actinal  spines  are  also  longer,  conical,  and  subacute. 

The  adambulacral  spines  are  slender  and  tapered,  becoming 
decidedly  longer  toward  the  mouth  and  on  the  adoral  plates.  The 
adoral  carinae  are  about  14  mm.  long,  strongly  compressed,  and  con- 
sist of  about  eleven  or  twelve  closely  united  pairs  of  adambulacral 
plates,  sloping  upward,  and  normally  bearing  each  one  long  spine; 
but  in  many  cases  one  is  lacking  in  a  pair,  and  on  some  of  the 
carinae  two  to  four  pairs  are  destitute  of  spines.  Those  in  the  last 
erect  pair  (epiorals)  on  the  jaw  are  longer  and  stouter  than  the 
others.  The  terminal  or  peroral  pairs  of  spines  are  horizontal,  short, 
stout,  flattened,  enlarged  or  subspatulate  at  the  blunt  tips.  A  short, 
thick,  strongly  lateral  divergent  spine,  not  half  as  long,  occupies 
the  side  angles. 

The  adambulacral  pores  are  large,  and  on  the  proximal  part  of  the 
rays,  within  the  radius  of  the  disk,  they  form  eight  distinct  rows; 
they  are  reduced  to  four  regular  rows  a  little  beyond  the  border  of 
the  disk.  On  about  three  adoral  pairs  of  plates,  they  also  decrease, 
the  innermost  being  a  single  pair. 

The  minor  pedicellariae  are  abundant  on  nearly  all  the  spines, 
but  the  wreaths  around  the  bases  of  the  dorsal  and  marginal  spines 
are  not  so  large  as  in  var.  australis.  Numerous,  large,  erect,  sessile, 
stout-ovoid  and  stone-hammer-shaped  pedicellariae,  with  strongly 
serrate,  blunt  jaws,  are  scattered  everywhere  over  the  back,  on  the 


86  VERRILL 

lateral  channels  and  among  the  ventral  spines.  Along  the  inner 
edges  of  the  adambulacral  grooves,  attached  by  pedicels,  there  are 
many  compressed  and  longer  ones,  some  of  which  are  larger  than 
the  dorsals,  others  much  smaller.  These  are  mostly  long-ovate, 
often  with  incurved  margins  and  blunt  at  the  tips,  which  are  den- 
tate ;  others  are  acute-lanceolate,  and  many  are  quite  small. 

In  the  Museum  of  Yale  University  there  are  also  several  smaller 
dry  specimens  of  this  species,  agreeing  very  closely  with  the  type. 

The  best  medium-sized  specimens,  with  the  radii  40  mm.  and 
180  mm.,  have  the  ratio  as  i :  4.5.  The  adambulacral  spines  are 
slender,  terete,  subacute,  arranged  in  one  simple  row.  Those  near 
the  mouth  are  not  much  lengthened;  more  distally  many  of  them 
are  somewhat  flattened,  with  obtuse  or  slightly  spatulate  tips.  They 
are  mostly  without  minor  pedicellariae,  though  small  groups  occur 
on  some  of  them.  A  few  bear  single,  rather  large,  ovate-lanceolate 
major  pedicellariae,  like  the  adambulacral  pedicellariae.  The  latter 
are  rather  numerous,  attached  to  the  inner  edge  of  the  adambulacral 
plates,  at  or  within  the  margin,  by  a  long  pedicel,  which  often  bears 
a  cluster  of  minute  acute  pedicellariae  at  and  below  the  base  of  the 
large  one ;  the  latter  is  unusually  large,  compressed-ovoid,  subacute 
or  obtuse,  with  the  tips  of  the  jaws  often  denticulate.  Their  pedicels 
are  often  as  long  as  the  adambulacral  spines.  The  actinal  spines 
are  large,  subequal,  and  form  four  or  five  regular  rows,  proximally, 
but  usually  only  four  beyond  the  middle  of  the  ray;  they  become 
much  more  slender  distally ;  those  of  the  interbrachial  region  become 
longer,  fusiform,  and  often  acute.  Most  of  them  are  stout,  clavate 
or  subcapitate,  strongly  sulcated  on  the  outside  and  with  the  tip 
finely  striated  all  around.  Most  of  them  bear  a  small  cluster  of 
minor  pedicellariae  on  the  outside.  There  are,  apparently,  only  four 
rows  of  peractinal  and  synactinal  plates.  The  outer  (peractinal) 
ones  often  bear  two  spines. 

The  marginal  spines  are  larger  and  rather  longer,  similar  to  the 
dorsal  spines,  and  with  subconic  striated  tips;  they  form  a  very 
regular  row,  standing  singly  and  well  spaced  on  a  row  of  somewhat 
raised  ossicles,  and  leave  below  them  a  wide  and  well  defined  mar- 
ginal channel,  which  becomes  dilated  at  the  interbrachial  region. 

The  dorsal  spines  are  similar  in  form  but  unequal  in  size,  numer- 
ous, arranged  without  any  very  evident  order,  but  pretty  evenly 
spaced.  They  mostly  stand  singly  on  the  nodes  of  the  reticulating 
ossicles,  which  form  five  to  seven  radial  ranges  and  nine  to  ten  in 
transverse  series.  The  dorsal  spines  bear  a  thin  basal  wreath  of 


SHALLOW-WATER  STARFISHES  87 

minor  pedicellariae.  There  are  also  many  small  clusters  of  dermal 
minor  pedicellariae.  Papulae  small,  very  numerous,  in  clusters. 

The  dorsal  major  pedicellariae  are  numerous,  scattered  among  the 
spines  and  especially  in  the  lateral  channels.  They  are  short,  thick, 
ovoid,  obtuse,  with  the  jaws  denticulated  at  the  apex;  they  often 
equal  the  smaller  spines  in  diameter,  but  are  shorter. 

This  species  can  only  be  confounded  with  P.  capitatus,  to  which  it 
is  evidently  closely  allied.  The  latter,  however,  not  only  has  much 
larger  and  more  capitate  dorsal  spines,  but  they  are  much  fewer  in 
number,  there  being  usually  only  about  five  or  six  spines  in  a  trans- 
verse series,  besides  the  laterals. 

The  supposed  absence  of  large  major  pedicellariae  on  the  back  of 
P.  Ititkenii,  referred  to  by  Stimpson,  does  not  hold  good,  as  shown 
by  our  specimens,  above  described. 

This  species  appears  to  range,  in  its  typical  form,  from  Van- 
couver Island  to  Monterey,  California,  and  as  a  variety  farther  south 
to  San  Diego. 

Oregon  (Stimpson)  ;  Saanich  Inlet,  Vancouver  Island  (de 
Loriol)  ;  San  Diego  and  Monterey,  California  (Yale  Museum)  ; 
Vancouver  Island  (Canadian  Geological  Survey)  ;  off  Pacific  Grove, 
near  Monterey,  taken  on  fish  lines  (Dr.  W.  R.  Coe.  Two  very  large 
specimens,  a,  b). 

When  full-grown  this  is  one  of  the  largest  starfishes  on  the  North- 
west Coast.  It  is  equalled,  in  this  genus,  only  by  P.  achraceus, 
P.  gigcmteus,  and  P.  papulosus. 

VARIATIONS. 

One  of  the  very  large  specimens  (&)  obtained  from  off  Monterey 
by  Dr.  Coe,  1901,  differs  considerably  from  the  more  typical  ones  in 
the  character  of  its  spinulation,  due  perhaps,  to  a  superabundance 
of  food. 

Its  radii  are  55  mm.  and  284  mm. ;  ratio,  about  1 : 5.2.  It  is 
noticeable  on  account  of  the  great  development  of  its  actinal  and 
inferomarginal  spines,  which  are  decidedly  more  numerous  and 
longer  than  in  the  other  specimen  of  equal  size  and  from  the  same 
place,  described  above. 

The  proximal  inferomarginal  and  most  of  the  actinal  plates  bear 
two,  three,  or  even  more,  stout,  often  divergent  spines,  so  that  one 
can  count  eight  to  ten  of  these  spines  in  each  irregular  transverse 
series.  But  one  ray  (doubtless  a  reproduced  one)  is  only  about  two- 
thirds  as  long  as  the  rest,  though  nearly  as  stout ;  on  this  the  ventral 


88  VERRILL 

spines  form  only  four  or  five  rows,  though  similar  in  size.  On  the 
distal  half  of  all  the  rays  the  ventral  rows  decrease  to  about  four  to 
six,  the  plates  bearing  only  two  spines,  or  but  one. 

The  ventral  spines  are  unusually  crowded  proximally.  They  are 
also  longer  and  less  stout,  though  mostly  blunt,  except  the  syn- 
actinals  and  those  on  and  near  the  interradial  areas,  where  many  of 
them  become  decidedly  longer  and  acute.  Many  in  the  synactinal 
row  are  deeply  grooved  on  the  outside,  and  the  distal  ones  become 
more  slender  and  subacute.  The  adoral  spines  are  long,  slender  and 
acute. 

Large  serrate  major  pedicellariae  are  scattered  over  the  dorsal 
and  lateral  surfaces,  but  are  not  half  so  numerous  as  on  the  other 
large  specimen. 

The  dorsal  spines  are  less  numerous,  larger,  more  acute,  and  more 
regularly  spaced,  their  average  intervals  being  8  mm.  to  10  mm. 
wide. 

Minor  pedicellariae  form  dense  wreaths  around  all  the  dorsal  and 
upper  marginal  spines. 

Off  Pacific  Grove,  California  (Dr.  W.  R.  Coe,  1901). 

PISASTER  LUTKENII  Var.  AUSTRALIS  Verrill,  nov. 

Two  very  large  specimens  (c,  d)  of  this  variety  are  in  the  Museum 
of  Yale  University.  One  has  the  radii  56  mm.  and  252  mm. ;  ratio, 
1:4.5. 

The  oral  adambulacral  spines  are  one-fourth  longer  than  those  of 
the  mid-ray ;  they  are  long,  slender,  regularly  tapered.  The  ventral 
spines,  near  the  base  of  the  rays,  are  in  four  and  five  crowded  rows ; 
but  on  the  mid-ray  there  are  about  four  regular  rows ;  those  of  the 
inner  and  middle  rows  are  very  stout  and  blunt,  often  nearly  as  thick 
as  long,  irregular  in  form,  often  pinched,  lobed,  or  sulcated,  and 
sometimes  slightly  bifid;  the  tips  are  often  clavate,  swollen,  or 
truncate. 

The  inferomarginals,  or  two  outer  rows,  and  those  near  the 
mouth,  are  longer,  not  so  stout,  and  more  regular,  mostly  with 
obtuse  or  ovoid  tips.  Distally  they  become  longer  and  more  fusi- 
form, and  many  are  deeply  channeled  on  the  outer  side. 

The  inferomarginal  spines  stand  two  on  a  plate,  nearly  or  quite 
to  the  tips  of  the  rays,  and  thus  form  the  two  outer  rows  of  ventral 
spines.  There  are  proximally  three  rows  of  stout  actinal  plates,  each 
usually  bearing  a  single  spine,  thus  forming  three  rows;  the  first 
row  (peractinal)  extends  to  the  tip  of  the  ray;  the  third  row  (syn- 
actinal) only  extends  to  about  the  distal  third  of  the  ray. 


SHALLOW-WATER   STARFISHES  89 

The  dorsal  spines  are  much  less  numerous  than  in  the  typical 
variety;  they  are  evenly  scattered,  and  scarcely  form  any  distinct 
radial  rows.  About  thirteen  to  fifteen  spines  occur  in  irregular 
transverse  series,  besides  the  upper  marginals,  but  they  do  not  form 
any  definite  transverse  rows. 

The  upper  marginals  are  much  like  the  dorsals,  but  larger.  They 
mostly  stand  singly  on  the  large  plates,  so  that  they  are  well  apart. 

Between  the  upper  and  lower  marginals  there  is  a  rather  wide, 
naked  channel,  bearing  large  pedicellariae;  at  the  base  of  the  rays  it 
expands  into  a  rather  large,  triangular,  interradial  area,  destitute 
of  spines. 

Minor  pedicellariae,  in  very  large  dense  wreaths,  surround  the 
upper  marginal  and  dorsal  spines,  the  clusters  becoming  so  large 
distally  that  they  are  in  contact.  In  dry  specimens  they  seem  to  be 
attached  outside  the  swollen  base  of  the  spines,  which  is  covered  by 
a  thick  skin,  but  in  life  they  probably  rise  up  at  least  to  mid-height 
of  the  spine.  On  the  ventral  spines  they  form  large  clusters  on  the 
outer  surface. 

Very  large,  ovoid,  or  blunt  stone-hammer-shaped,  erect,  sessile, 
major  pedicellariae,  with  strongly  serrate  jaws,  occur  in  considerable 
numbers  on  the  lateral  channels,  between  the  dorsal  spines,  and 
among  the  ventral  spines.  Other  major  pedicellariae,  some  of  them 
nearly  as  large,  but  others  much  smaller,  somewhat  compressed, 
mostly  ovate  and  obtuse,  occur  on  the  adambulacral  spines  and  on 
the  inner  edge  of  the  ambulacral  grooves.  Many  of  these  have  large 
numbers  of  small,  acute,  forficulate  pedicellariae  on  their  pedicels. 

San  Diego,  California  (Dr.  Edw.  Palmer,  Yale  Museum). 


PISASTER  GIGANTEUS  (Stimpson)  Verrill. 
Plate  xxxvn,  figures  i,  2  (type). 

Asterias  gigantea  STIMPSON,  Jour.  Boston  Soc.  Nat.  Hist.,  vi,  p.  88,  pi.  xxni, 
figs.  4-6,  1857.  Verrill,  Trans.  Conn.  Acad.,  i,  p.  327,  1867.  ?  Bell  (pars), 
Proc.  Zool.  Soc.  London,  1881,  p.  564;=:  in  part  A.  katherina  Per.,  non 
Gray. 

Dr.  Stimpson's  incomplete  original  description  is  as  follows: 
"  Body  very  large,  swollen  ;  rays  six  in  number,  in  length  some- 
what less  than  twice  the  diameter  of  the  disk.  Upper  surface  cov- 
ered with  numerous  short,  blunt,  equidistant  spines,  uniform  in  size 
and  regularly  distributed;  these  spines  are  somewhat  conical  in 
shape,  but  truncated  at  the  tip  and  constricted  at  the  base,  with  the 
sides  longitudinally  furrowed.  The  spines  of  the  lower  surface 


90  VERRILL 

are  short  and  thick,  but  slightly  compressed  and  notched  at  the 
extremity.  Diameter,  two  feet.  Taken  in  Tomales  Bay,  by  Mr. 
Samuels." 

Large  photographs  of  the  type,  which  is  in  the  U.  S.  National 
Museum,  have  been  furnished  by  Dr.  R.  Rathbun,  for  reproduction 
in  our  plates.  Dr.  Stimpson's  description  was  very  incomplete,  for 
he  scarcely  described  the  ventral  side  at  all. 

The  disk  is  rather  large  and  the  six  rays  taper  rapidly  to  rather 
acute  tips.  They  are  variously  bent,  and  wrinkled,  indicating  a 
rather  weakly  reticulated  dorsal  skeleton.  The  radii  are  about 
73  mm.  and  304  mm. ;  ratio,  about  1 : 4.75. 

The  adambulacral  spines  are  rather  small,  slender,  compressed, 
crowded,  and  stand  in  a  single  row,  one  to  a  plate;  they  are  much 
smaller  and  shorter  than  the  crowded  ventral  spines.  The  latter 
form  about  eight  close  rows  proximally,  but  these  decrease  to  four 
or  five  at  the  distal  third  of  the  rays.  They  differ  much  in  form. 
Those  nearer  the  adambulacrals  are  partly  fusiform  and  sometimes 
subacute,  but  more  often  flattened  and  subacute  at  the  tip;  those  of 
the  middle  rows  are  stouter,  either  blunt,  clavate,  or  flattened ;  those 
of  the  outer  rows  are  still  shorter  and  stouter,  with  blunt  tips,  but 
those  near  the  base  of  the  rays  are  apt  to  be  fusiform  and  subacute. 

The  lateral  or  superomarginal  row  of  spines  is  not  very  distinct; 
its  spines  are  subacute  and  much  smaller  than  the  actinals.  They 
form  in  some  places  two  irregularly  alternating  rows,  and  differ  but 
little,  in  size  or  form,  from  the  dorsal  spines,  though  they  may  be 
rather  longer  and  more  acute. 

The  dorsal  spines  are  small,  either  obtuse  or  acute,  roughly 
striated,  very  numerous,  nearly  uniformly  scattered  over  the  whole 
surface,  mostly  standing  singly,  and  do  not  form  reticulations  nor 
any  distinct  median  row.  On  the  sides  of  the  rays,  in  some  places, 
they  form  small  transverse  groups  or  combs  on  the  connective 
ossicles.  About  twenty  to  twenty-four  can  be  counted  in  an  irregular 
series  across  the  rays  at  the  base,  but  they  form  no  evident  trans- 
verse rows. 

The  spines  become  rather  larger,  longer,  and  more  crowded  near 
the  ends  of  the  rays,  where  they  are  surrounded  by  close  wreaths  of 
minor  pedicellariae.  On  the  basal  part  of  the  rays  and  on  the  disk 
the  minor  pedicellariae  form  small  clusters  around  the  bases  of  the 
spines,  and  many  other  dermal  ones  are  scattered  between  the  spines. 

A  few  rather  small,  stout,  subconical,  dermal  major  pedicellariae 
are  scattered  on  the  dorsal  and  lateral  surfaces,  and  especially  on  the 


SHALLOW- WATER   STARFISHES  91 

interbrachial  areas  and  intermarginal  channels.  The  actinal  spines 
bear  small  clusters  of  minor  pedicellariae  on  their  outer  sides. 

This  species  appears  to  be  very  distinct  from  all  others.  Perrier 
placed  it  as  a  synonym  of  his  A.  katherin&=P.  grayi  Ver.,  to  which 
it  bears  some  little  resemblance  in  form.  He  was  probably  misled  by 
Stimpson's  very  poor  description. 

Nothing  is  known  concerning  its  range.  Stimpson's  original 
record  is  the  only  authentic  locality  known  to  me.  Some  of  the 
localities  recorded  are  due  to  errors. 

PISASTER  PAPULOSUS  Verrill. 

Plate  XLII,  figure  i;  plate  XLIII,  figure  i;  plate  LX,  figure  i;  plate  LXXVI, 

figures  2-2d  (details)  ;  plate  LXXX,  figure  4. 
Pisaster  papulosus  VERRILL,  Amer.  Journ.  Sci.,  vol.  xxvin,  p.  63,  1909. 

The  type  of  this  species  is  a  large  dry  five-rayed  specimen,  in 
excellent  condition,  taken  in  Puget  Sound  and  sent  to  me  by  Pro- 
fessor T.  Kincaid.  A  specimen  received  from  the  Provincial 
Museum  of  British  Columbia,  through  Dr.  C.  F.  Newcombe,  but  not 
in  so  good  preservation,  is  much  larger  (diameter,  28  inches  or 
660  mm.).  Thus,  when  full-grown  it  is  one  of  the  largest  starfishes 
yet  discovered. 

The  disk  is  large  and  remarkably  high  and  plump,  with  a  raised 
central  pentagon,  bearing  ten  groups  of  spines  on  the  angles  and 
sides,  besides  a  central  group ;  from  the  radial  angles  carinal  radial 
rows  of  spines  run  to  the  tips  of  the  arms,  which  are  long,  rather 
stout  at  the  base,  but  regularly  tapered  to  acute  tips. 

The  radii  of  the  type  are  42  mm.  and  215  mm.  to  225  mm. ;  ratio, 
about  i :  5.25 ;  breadth  of  rays  at  base,  45  mm. ;  elevation  of  disk, 
46  mm.;  length  of  dorsal  spines,  4  mm.;  diameter,  i  mm.;  length 
of  larger  ventral  spines,  3  mm.  to  4  mm. ;  diameter,  2  mm.  to  2.5  mm. 

The  dorsal  surface  is  covered  with  an  integument  that  conceals  the 
ossicles.  The  papular  areas  are  large  and  numerous ;  a  conspicuous 
row  runs  along  each  side  of  the  median  radial  row  of  spines,  and 
there  are  several  less  regular  rows  on  each  side.  The  papulae  are 
very  numerous  and  small,  darker  than  the  surrounding  integument, 
which  is  light  yellowish  orange  in  the  dried  specimen;  the  papular 
areas  brown. 

The  dorsal  spines  are  large,  few,  and  mostly  isolated,  but  they 
form  a  distinct,  simple,  radial  or  carinal  row,  one  to  a  plate,  proxi- 
mally,  but  on  the  distal  third  of  the  ray  these  spines  become  more 
distant  and  the  row  less  distinct. 


92  VERRILL 

On  either  dorso-lateral  side  there  are  similar,  but  rather  sparsely 
scattered,  spines,  which  mostly  show  no  definite  arrangement ;  but  in 
some  cases,  distally,  they  have  a  tendency  to  form  about  three 
imperfect  rows  on  each  side.  These  spines  are  all  rather  large,  up  to 
3  mm.  to  4  mm.  high,  and  about  I  mm.  thick,  tapered  or  conical, 
subacute  or  acuminate,  and  often  pinched  up  near  the  tips  and 
strongly  sulcated.  They  bear  large  and  dense  wreaths  of  minor 
pedicellariae  placed  around  their  bases  as  dried,  and  there  are  numer- 
ous large,  dense  clusters  of  similar  pedicellariae  thickly  scattered 
over  the  surface  between  the  spines,  and  around  the  papular  areas, 
as  if  taking  the  places  of  the  spines  that  are  present  in  other  species. 
Many  of  these  clusters  are  as  large  as  those  around  the  spines,  or  up 
to  3  mm.  to  4  mm.  in  breadth.  They  are  abundant  on  both  the  disk 
and  rays,  but  become  more  numerous  on  the  distal  half  of  the  rays, 
where  they  are  often  so  crowded  that  in  life  they  must  form  a 
nearly  continuous  coating.  Large,  stout,  erect,  dentate,  dermal, 
major  pedicellariae  are  numerous,  scattered  over  the  whole  dorsal  and 
lateral  surfaces  of  the  rays.  They  are  rather  quadrangular  in  an 
end  view;  in  a  side  view  scarcely  compressed,  blunt  wedge-shaped 
or  stone-hammer-shaped.  The  tip  of  the  jaws  are  serrate  or 
unguiculate  and  subtruncate.  The  larger  ones  are  about  1.5  mm. 
high  and  nearly  i  mm.  broad. 

The  pentagon  of  the  disk  is  formed  by  five  radial  clusters  of  stout 
conical  spines,  like  the  dorsal  radials,  standing  about  three  together, 
and  five  similar  interradial  groups,  standing  a  little  nearer  the  center. 
The  large,  brown  madreporic  plate  takes  the  place  of  most  of  those  in 
one  interradial  space.  A  cluster  of  five  spines  occupies  the  center. 

The  large  dorsal  interradial  areas  of  the  disk  are  mostly  without 
spines,  but  have  an  abundance  of  pedicellariae  and  papulae. 

The  upper  marginal  rows  of  spines  are  like  the  median,  but  they 
become  more  regular  and  the  spines  closer  together  distally.  The 
channel  below  them  is  wide  proximally,  and  bears  many  of  the  large, 
serrate  pedicellariae  and  numerous  minor  ones.  The  row  of  infero- 
marginal  spines  is  strong,  double,  and  regular,  each  plate  usually 
bearing  two  spines,  especially  distally,  to  the  tips  of  the  rays.  The 
distal  spines  are  subacute  and  sulcate,  much  like  the  dorsals,  but 
those  on  the  proximal  half  are  mostly  stouter  (2  mm.  or  more),  with 
obtusely  rounded  and  sulcated  tips. 

Close  to  these,  but  with  an  intervening  papulose  channel,  is  a 
regular,  mostly  single  row,  of  peractinal  spines,  continuous  to  the 
tips  of  the  rays.  The  spines  are  like  the  inferomarginals.  A 


SHALLOW-WATER   STARFISHES  93 

shorter  row  of  similar  synactinal  spines,  extending  to  the  distal 
third,  intervenes  between  the  peractinals  and  the  adambulacrals,  but 
is  separated  from  the  latter  by  a  definite  channel,  having  papulae 
proximally.  Close  to  the  base  of  the  rays  some  of  the  infero- 
marginal  and  actinal  plates  often  bear  two  or  three  small  extra 
spines,  and  all  bear  dense  clusters  of  minor  pedicellariae  on  the  outer 
side.  The  adambulacral  spines  form  a  single  very  regular  row; 
they  are  slender,  tapered,  and  subacute;  about  five  correspond  to  a 
single  actinal  spine.  The  adoral  ones  become  somewhat  longer. 
Many  of  them  bear  thick  clusters  of  forficulate  pedicellariae  of 
various  sizes  in  a  single  cluster.  Between  their  bases  and  on  the 
inner  edges  of  the  ambulacral  grooves,  and  on  the  oral  spines,  and 
interradial  areas  are  many  large  major  pedicellariae,  part  of  which 
are  unguiculate,  like  those  of  the  back,  but  others  are  longer,  com- 
pressed, lanceolate,  with  the  tips  subacute,  or  terminating  in  one  or 
two  hook-like  teeth.  The  pedicels  of  these,  and  also  of  those 
attached  to  the  spines,  are  often  surrounded  by  large  clusters  of 
small  forficulate  pedicellariae  of  nearly  the  same  form,  but  of  various 
sizes,  many  being  as  minute  as  the  ordinary  minor  pedicellariae. 
Friday  Harbor,  Puget  Sound  (Professor  Kincaid). 

The  paucity  of  spines  on  the  back  of  the  specimen  above 
described  might  be  thought  to  be  due  in  part  to  some  injury  and 
lack  of  subsequent  restoration  of  the  spines.  But  on  the  distal  part 
of  the  rays,  at  least,  the  spines  seem  to  be  certainly  normally 
arranged.  There  are  no  naked  ossicles  or  tubercles  where  spines 
are  likely  to  have  been  attached. 

I  have  examined  four  additional  specimens  (No.  1904,  Mus. 
Comp.  Zool.)  from  the  same  district  and  also  sent  by  Professor 
Kincaid.  They  are  all  larger.  One  of  them  agrees  closely  with  the 
type  specimen  described  above.  Its  radii  are  1 10  mm.  and  330  mm. 
Others,  equally  large,  have  the  disk  and  rays  flattened  in  drying 
and  appear  to  have  more  numerous  and  larger  spines  and  fewer 
pedicellariae.  One  has  the  radii  85  mm.  and  330  mm. 

This  species  is  rather  closely  related  to  P.  brevispinus  (Stimpson). 
But  the  characters  of  the  dorsal  spines  and  pedicellariae  and  of  the 
actinal  spines,  and  the  larger  spines  and  papular  areas,  seem  to 
require  their  separation.  Of  course  a  much  larger  series  of  specimens 
than  I  have  studied  might  show  intermediate  forms,  as  in  other  cases. 
The  difference  in  the  number  of  rows  of  actinal  spines  is  greater  than 
we  should  expect  in  specimens  of  similar  size.  The  dorsal  spines 
are  much  less  numerous  in  our  species  and  the  rays  much  longer 


94  VERRILL 

and  more  tapering.    The  characters  of  the  adoral  carina  and  oral 
spines  are  also  peculiar. 

VARIATIONS. 

A  very  large  dry  specimen  of  this  species  has  been  sent  to  me  by 
Dr.  C.  F.  Newcombe  from  the  Provincial  Museum  of  British 
Columbia.  It  was  dredged  in  ten  fathoms,  off  Victoria,  1894. 

Its  radii  are  64  mm.  and  330  mm. ;  ratio,  1 :  5.14. 

The  disk  is  distorted  in  drying  and  is  probably  larger  than  the 
normal  size.  The  rays  are  thick  at  base,  but  taper  to  long  and 
relatively  slender  tips.  The  dorsal  surface  is  covered  with  a  thick 
integument,  dark  reddish  brown  in  the  dry  specimen,  and  has  but  few 
distant  spines.  But  numerous  large,  stout,  thick,  unguiculate  pedi- 
cellariae  are  scattered  over  the  surface,  which  is  also  covered  with 
small  minor  pedicellariae,  thickly  scattered  over  nearly  the  whole 
surface  and  also  forming  wreaths  around  the  bases  of  the  spines. 
Probably  a  large  part  of  these  may  have  been  rubbed  off  by  repeated 
dustings,  for  the  specimen  has  evidently  been  exposed  to  dust  and 
mold;  and  owing  to  its  great  size,  it  probably  was  not  very  well 
preserved  originally.  Hence  it  is  probable  that  the  dermal  minor 
pedicellariae  originally  existed  in  still  larger  clusters,  as  they  do  in 
depressed  spots,  where  the  coating  of  dust  and  mold  has  not  been 
removed,  especially  distally.  But  as  numerous  large  major  pedi- 
cellariae are  still  scattered  over  the  surface,  it  is  evident  that  no 
severe  cleansing  has  taken  place,  sufficient  to  remove  all  those  on 
the  spines. 

The  dorsal  skeleton  is  coarsely  and  irregularly  reticulated,  but 
the  ossicles  are  concealed  by  the  thick  integument. 

The  spines  are  mostly  conical  and  acute  or  subacute,  about  1.5  mm. 
to  2.5  mm.  long,  with  but  few  smaller  ones.  They  form  distinct  but 
incomplete  simple  radial  rows,  in  which  they  usually  stand  2  mm.  to 
5  mm.  apart,  where  continuous.  A  very  few  similar  spines  are 
irregularly  scattered  over  the  disk  and  rays.  The  madreporic  plate 
is  large,  with  fine  gyri,  its  diameter  is  13  mm. 

The  superodorsal  plates  are  well  down  on  the  sides  of  the  rays. 
They  are  relatively  small  and  not  very  distinct,  and  bear  a  simple 
row  of  spines,  rather  larger  than  the  dorsals,  many  of  them  blunt 
with  grooved  tips.  They  are  close  to  the  inferomarginals,  except 
in  the  interradial  areas,  where  they  diverge  a  little,  leaving  a  spine- 
less area. 

The  ventral  spines  proximally  consist  of  four  or  five  rows  close 
together  and  nearly  equal.  They  are  short  (3  mm.  to  4  mm.  long), 


SHALLOW-WATER   STARFISHES  95 

stout,  mostly  blunt  or  truncate  and  striated  at  the  tip ;  but  close  to 
the  bases  of  the  rays  they  become  longer.  Some  are  acuminate  and 
subacute.  On  the  distal  third  of  the  rays  there  are  only  three  rows, 
besides  the  superomarginals,  there  being  but  one  row  of  actinals 
distally. 

The  inferomarginal  ossicles  are  rather  large,  thick  and  convex, 
roundish  on  the  outer  end,  and  usually  bear  two  or  three  divergent 
spines,  which  are  mounted  on  low  mammillae  with  a  central  pit.  The 
peractinals  are  similar  in  form  externally,  and  mostly  bear  but  one 
spine,  but  sometimes  two  proximally.  The  principal  row  of  sub- 
actinal  plates,  proximally,  are  large  and  transversely  broad  in  their 
internal  part,  but  show  only  a  rounded  exterior  surface,  smaller 
than  the  other  plates  and  bearing  a  single  spine.  Between  these, 
proximally,  there  are  a  few  smaller  wedge-shaped  or  irregular  sub- 
actinal  ossicles,  interpolated  irregularly  and  mostly  without  spines. 
The  inferomarginal  and  actinal  plates  are  closely  crowded  together. 

There  is  a  rather  wide  naked  channel  between  the  synactinal  and 
adambulacral  plates,  due  to  the  great  transverse  length  of  the  latter 
externally  to  the  single  spine  that  stands  on  the  inner  edge. 

The  adambulacral  spines  are  slender  and  tapered,  mostly  5  mm. 
to  7  mm.  long,  becoming  still  longer  near  the  mouth. 

Large  major  pedicellariae,  like  those  of  the  type,  occur  among  the 
actinal  adambulacral  spines  and  along  the  inner  edge  of  the  grooves. 
These  are  long-ovate  or  lanceolate,  with  acute  or  subacute  jaws, 
often  toothed.  With  these  there  were  others  of  various  smaller 
sizes,  of  nearly  the  same  forms. 

THE  YOUNG. 

A  small  specimen,  from  Sitka,  has  the  radii  10  mm.  and  45  mm. ; 
ratio,  i :  4.5. 

The  dorsal  spines  are  few  and  sparse,  except  along  the  median 
radial  rows,  where  they  form  pretty  close  and  nearly  regular  simple 
rows.  A  very  imperfect,  irregular  row  also  occurs  on  each  side, 
about  midway  between  the  median  and  superomarginal  rows.  Ten 
small  groups  of  about  two  or  three  spines  each  form  a  central 
pentagon,  with  three  central  spines.  Elsewhere  dorsal  spines  are 
mostly  lacking.  All  these  spines  are  short,  cylindrical  or  slightly 
clavate,  with  obtuse,  sulcated  tips.  The  dorsal  ossicles  are  rather 
large  and  broad;  even  those  that  are  without  spines  are  often 
lozenge-shaped;  those  that  bear  spines  are  convex  centrally,  with  a 
mammilla  and  pit.  The  superomarginal  spines  form  a  regular  simple 


96  VERRILL 

row ;  they  are  smaller  than  the  dorsals,  but  of  the  same  form.  The 
inferomarginal  spines  stand  mostly  two  on  a  plate  and  with  the  per- 
actinals  form  three  regular  nearly  equal  rows.  They  are  larger  than 
the  superomarginals  and  either  cylindrical  or  slightly  tapered, 
obtuse,  finely  sulcate.  The  peractinals  are  rather  smaller  than  the 
rest  and  less  obtuse. 

Adambulacrals  form  a  single  row;  they  are  about  as  long  as  the 
actinals,  but  much  more  slender  and  evenly  tapered.  Three  or  four 
of  the  inner  adoral  plates,  forming  the  compressed  carinae,  are  with- 
out spines. 

A  few  large,  ovate,  unguiculate  pedicellariae  occur  on  the  lateral 
channels  and  adambulcral  plates.  Some  smaller  acute-lanceolate 
ones  occur  on  the  inner  edges  of  the  grooves.  Minor  pedicellariae 
are  rather  thickly  scattered  everywhere  over  the  dorsal  and  lateral 
integument. 

Another  young  specimen  of  larger  size,  with  the  radii  16  mm.  and 
82  mm.,  was  sent  by  Dr.  C.  F.  Newcombe.  It  was  taken  at  Kuper 
Island,  B.  C.,  February,  1894.  This  agrees  closely  with  the  smaller 
one,  described  above,  in  the  number  and  character  of  its  dorsal 
spines,  but  the  ventral  spines  are  relatively  stouter  and  form  four 
rows  proximally. 

The  dorsal  minor  pedicellariae  are  much  more  numerous,  both  on 
the  integument  and  around  the  spines,  especially  distally,  where  they 
form  large  dermal  clusters.  The  papular  areas  are  also  very  large, 
with  numerous  papulae.  Thus  it  has  in  most  respects  the  characters 
of  the  adult. 

Large  major  pedicellariae  are  rather  numerous  on  the  interradial 
areas  and  sides  of  the  rays ;  and  many  large,  very  acute,  lanceolate 
ones  occur  on  the  inner  edges  of  the  grooves,  sometimes  in  regular 
rows,  one  to  a  plate,  for  some  distance.  Most  of  these  are  sur- 
rounded at  base  by  clusters  of  smaller  major  pedicellariae  of  about 
the  same  shape,  but  of  various  sizes. 

This  specimen  is  remarkable  for  having  two  spines  on  some  of  the 
adambulacral  plates,  here  and  there,  the  second  spine  being  smaller 
and  attached  back  of  the  regular  one.  This  condition  has  not  been 
noticed  in  the  other  specimens. 

The  adoral  carinse  are  rather  long  and  strongly  compressed,  de- 
scending abruptly  to  the  mouth  in  all  our  specimens,  and  destitute  of 
spines  on  three  or  four  adoral  plates.  Apical  peroral  spines  long, 
tapered,  about  equal  to  adorals,  side-spines  apparently  lacking,  or 
replaced  by  a  cluster  of  pedicellariae;  epioral  spines  lacking  on  our 
specimens. 


SHALLOW-WATER  STARFISHES  97 

This  species  ranges  from  Puget  Sound  to  Sitka.  Puget  Sound 
(Professor  Kincaid) ;  Kuper  Island  and  Victoria  (C.  F.  New- 
combe)  ;  Sitka  (Harriman  Expedition,  W.  R.  Coe). 

PISASTER  GRAYI  Verrill,  nom.  nov. 

Asterias  katherince  PERKIER,  Steller.  du  Mus.,  Archiv  Zool.  Exper.,  iv,  p.  331, 

1875  (non  Gray,  1840). 
Asterias  dubia  VERRILL,  Amer.  Naturalist,  xun,  p.  545,  1909  (non  Clark). 

Perrier  evidently  erred  in  his  determination  of  Gray's  type  of 
A.  katherincz,  for  Gray  stated  that  his  species  had  two  spines  on  each 
adambulacral  plate,  as  I  have  elsewhere  explained.1 

According  to  Perrier  it  has  the  following  characters : 

Ordinarily  six  rays,  sometimes  five,  upper  side  of  the  body  little 
convex,  rays  broad  at  the  base,  pointed  at  the  tip.  Radii  as  1:7. 
Between  tips  of  rays  about  4  deem.  Adambulacral  spines  in  a  single 
range,*  with  some  small  clusters  of  major  pedicellariae  upon  the  outer 
side.  Immediately  beyond  these  comes  a  band  of  stouter  ventral 
spines,  formed  by  transverse  ranges  of  two  or  even  three  spines. 
A  narrow  channel  separates  this  band  from  another  simple  range  of 
spines,  representing  the  lateral  spines.  Then  comes  a  pretty  wide 
naked  band,  and  a  very  irregular  range  of  shorter  spines,  which 
indicates  the  commencement  of  the  dorsal  region.  The  latter  is  cov- 
ered with  numerous  short  spines,  with  the  head  rounded  and  strongly 
striated;  sometimes  isolated,  sometimes  in  groups  of  two  or  three, 
but  disposed  without  order.  A  circle  of  minor  pedicellariae,  small  in 
number,  surrounds  the  base  of  these  spines.  Some  small  major  pedi- 
cellariae are  scattered  between  them,  but  these  are  particularly 
numerous  on  the  sides,  between  the  back  and  and  the  line  of  lateral 
spines,  in  the  channel  that  separates  this  line  from  the  band  of 
ventral  spines,  and  between  the  latter  and  the  adambulacral  spines. 
These  major  pedicellariae  have  the  form  of  a  short  isosceles  triangle 
with  a  pretty  large  base.  The  madreporic  plate  is  marginal.  Color, 
when  dry,  red. 

Gray  gave  the  locality  of  his  specimens  as  "  Mouth  of  the  Colum- 
bia." This  locality  is  uncertain,  however;  for  his  specimens,  as 
labelled,  were  a  mixed  lot.  (See  p.  113,  below.) 

Perrier  regarded  A.  gigantea  Stimpson  as  a  doubtful  synonym  of 
this  species,  to  which  it  is  possibly  related.  But  gigantea  has  a  much 

1  Gray's  true  type  is  doubtful.    (See  page  113.)    It  could  not  have  been  this 
species. 
"This  does  not  agree  with  Gray's  statement.    (See  below,  p.  112.) 

8 


98  VERRILL 

larger  disk  and  shorter  rays.  The  latter  also  has  more  numerous 
rows  of  actinal  spines,  while  the  dorsal  spines  are  conical,  numer- 
ous, and  evenly  scattered.  Nor  does  Perrier  mention  the  large  ser- 
rate, lateral  pedicellariae  found  on  the  latter.1  Still  it  may  prove  to 
be  the  young  of  P.  giganteus,  for  we  know  very  little  about  the 
variations  of  the  latter. 

I  have  seen  no  specimens  agreeing  well  with  the  description  given 
by  Perrier. 

PISASTER  ?  PAUCISPINUS  (Stimpson)  Verrill. 
Plate  xxxvi,  figures  i,  2  (types). 

Asterias  paucispina  STIMPSON,  Proc.  Boston  Soc.  Nat.  History,  vm,  p.  266, 
1862.  Perrier,  op.  cit.,  1875,  p.  324  (no  description).  Bell,  op.  cit.,  p.  495 
(no  description). 

This  species  is  easily  recognized  by  its  rather  angular  rays,  few 
actinal  and  dorsal  spines,  and  uniserial  adambulacral  spines. 
Stimpson's  description  is  as  follows : 

"  Rays  five,  high,  trigonal  above,  rounded  below ;  disc  rather  large. 
Proportion  of  the  diameters,  1 : 4.75.  Skin-skeleton  less  firm  than 
in  the  four  preceding  species,  the  net-work  being  more  open.  Ambu- 
lacral  pores  in  four  regular  rows.  Ambulacral  spines  in  one  very 
regular  row,  equal,  not  crowded,  slightly  compressed,  and  slightly 
tapering,  with  blunt  extremity.  A  considerable  number  of  sharp, 
appressed  major  pedicellariae,  of  variable  size,  may  be  seen  on  the 
inter-ambulacral  plates  at  the  inner  bases  of  the  ambulacral  spines, 
together  with  a  few  small  clusters  of  the  minor  kind.  Ventral 
spines  of  moderate  size,  cylindrical,  tapering  to  a  blunt  tip  not 
striated,  and  arranged  in  three  rows,  or  in  about  thirty  transverse 
rows  of  three  each,  the  two  outer  ones  placed  together  on  each  of 
the  ossicles  of  the  single  ventral  [inferomarginal]  series,  and  the 
inner  one  on  the  transverse  connective  piece  [actinal  plate]  which 
passes  to  the  marginal  interambulacral  plates.  Each  of  these  trans- 
verse rows  corresponds  to  five  ambulacral  spines.  A  small  cluster 
of  minor  pedicellarise  at  the  outer  base  of  each  of  the  ventral  spines, 
most  prominent  in  those  of  the  outer  row.  Lateral  channel  distinct, 
with  a  row  of  stout,  narrow,  wedge-shaped  major  pedicellariae, 
extending  from  the  base  of  the  ray  to  the  middle  of  its  length. 
Dorsal  spines  equalling  the  ventrals  in  size,  less  than  one-twelfth 
inch  in  height,  and  subcapitate,  with  conical,  truncate,  and  striated 

1  If  these  were  absent  the  species  may  not  be  closely  related  to  the  Pisaster 
group,  especially  as  there  are  few  actinal  rows  of  spines. 


SHALLOW-WATER  STARFISHES  99 

heads.  They  are  few  in  number,  and  are  arranged  in  five  regular 
rows;  those  of  the  marginal  row,  twenty-five  in  number,  being  a 
little  smaller  and  more  elongated  than  the  others  ;  median  row  con- 
sisting, like  the  marginal  row,  of  twenty-five  spines,  one  to  each 
ossicle;  intermediate  row  with  only  ten  spines,  and  becoming 
"  zigzag  "  toward  the  extremity  of  the  ray.  On  the  disc  there  is 
a  regular  pentagon  of  about  ten  spines,  one  (rarely  two)  to  each 
angle,  and  one  (rarely  two)  at  the  middle  of  each  of  the  concave 
sides.  No  spines  within  the  pentagon  except  one  central  one,  which 
is  always  present;  madreporic  plate  within  the  pentagon,  at  the 
periphery.  Minor  pedicellarise  are  scattered,  in  clouds,  over  the 
dorsal  surface,  but  there  are  more  of  the  major  kind  on  the  back. 
Papulae  in  groups.  Diameter,  four  and  a  half  inches. 

"  Habitat,  Puget  Sounds-North  West  Boundary  Commission. 
Dr.  C.  B.  Kennerlyv—  ^Fhts  fine  species  is  common  in  the  circumlit- 
toral  zone." 


The  figures  are  from  photographs  of  Stimpson's  original  types  in    & 
the  U.  S.  National  Museum.    They  were  furnished  by  Dr.  Richard 

T>     4.UU  TU  •  U     1.1        • 

Rathbun.    These  specimens  are  probably  immature.  ^.      _-..£*, 


VARIATIONS. 

A  purchased  specimen  in  the  Museum  of  Yale  University,  labelled 
simply  as  from  the  "  West  Coast,"  agrees  closely  with  Stimpson's 
type,  except  in  characters  due  to  its  smaller  size.  Its  radii  are 
10  mm.  and  39  mm.  ;  ratio,  about  1  :  4.  The  rays  are  high  and 
somewhat  carinate.  The  dorsal  ossicles  are  not  numerous,  rather 
stout  and  broad,  but  have  rather  broad  rounded  papular  areas;  the 
ossicles  of  the  median  row  are  small  but  prominent,  each  bearing, 
near  the  base  of  the  arm,  two  short  obtuse  or  subcapitate  spines,  but 
distally  only  one.  Between  the  median  and  the  superolateral  row 
there  is  a  single  row  of  fewer,  larger,  obtuse  spines,  with  striated 
ends.  The  superomarginals  are  a  little  longer  and  somewhat  conical, 
one  to  a  plate.  The  pentagon  of  spines  on  the  disk  has  usually  two  or 
three  spines  at  each  angle,  one  or  two  on  each  side,  and  two  on  one 
ossicle  in  the  middle  ;  all  these  are  capitate  and  striate.  Madreporic 
plate  raised,  wart-like,  with  fine  radiating  gyri.  The  under  side  is 
nearly  as  described  by  Stimpson. 

The  adambulacral  spines  are  uniserial,  very  slender,  terete,  tapered, 
subacute.  The  synactinal  plates  are  small,  rounded,  and  prominent, 
extending  to  about  the  distal  fourth  of  the  ray,  each  bearing  a  spine 
like  the  inferomarginals,  but  rather  smaller.  The  latter,  which 


IOO  VERRILL 

mostly  stand  two  to  a  plate,  are  tapered,  subacute  or  obtuse,  but 
not  striated  at  tip. 

Large  major  pedicellarise  occur  in  the  naked  channels  between  the 
upper  and  lower  marginal  plates,  especially  proximally;  these  are 
compressed  in  a  dorsal  view,  but  long-ovate  or  lanceolate  in  a  side 
view,  not  serrate.  These  are  more  than  half  as  thick  as  the  adjacent 
spines ;  others  of  much  smaller  size,  but  similar  in  form,  occur  on  the 
interradial  areas  below,  and  along  the  margin  of  the  grooves.  The 
minor  pedicellariae  are  very  small  and  occur  both  on  the  spines  and 
on  the  papular  areas. 

This  species  may  belong  with  Marthasterias  rather  than  with 
Pisaster.  More  specimens  are  needed.  Perhaps  larger  specimens 
would  have  more  rows  of  interactinal  plates. 

Genus  Marthasterias  Jullien. 

Asterias  (Pars)  GRAY,  Sladen,  Bell,  etc. 

Marthasterias  JULLIEN,   Bull.  Zool.   Soc.   France,   1878,  p.   141.     Type,  M. 

foliacea  =  M .  glad  alts  (Mull.). 

Stolasterias,  subgenus  (pars),  SLADEN,  op.  cit,  1889,  pp.  563,  583. 
Stolasterias  (restr.)  PERKIER,  Exped.  Trav.  et  Talisra.,  pp.  108,  109,  1894. 

Monacanthid,  five-  or  six-rayed  Asteriinae,  with  few  definite  radial 
rows  (usually  three)  of  stout  dorsal  ossicles  and  large  spines.  One 
row  of  small  or  rudimentary  actinal  plates,  usually  without  spines. 
Upper  and  lower  marginal  plates  large,  with  long  spines.  See  also 
P-  47- 

MARTHASTERIAS    (?)    SERTULIFERA   (Xantus)   Verrill. 

Asterias  sertulifera  XANTUS,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  1860,  p.  568. 
Ives,  List  Cab.  Stearns,  p.  2,  1890.  De  Loriol,  Recueil  Zool.,  Suisse,  n, 
1887. 

Rays  five,  subangular.  Radii,  about  1:5;  diameter,  about  112  mm. 
Adambulacral  spines  in  one  row,  very  slender,  flattened.  Ventral 
spines  in  two  or  three  rows,  stouter,  blunt,  and  flattened.  Upper 
marginals  regular,  one  spine  to  each  alternate  plate.  Dorsal  spines 
similar,  stout,  cylindrical  or  tapered,  in  three  pretty  regular  rows,  all 
bearing  dense  wreaths  of  minor  pedicellariae  near  their  tips.  Major 
pedicellariae  few.  (Xantus.) 

Cape  St.  Lucas  (Xantus)  ;  San  Diego  (Ives,  1890)  ;  California 
(De  Loriol). 

I  have  had  no  opportunity  to  study  well  preserved  examples  of  this 
species,  and  therefore  refer  it  to  Marthasterias  with  doubt. 


SHALLOW-WATER   STARFISHES  IOI 

Genus  Asterias  Linne  (restricted).    See  page  43. 

^Asterias  (pars)  LINNE,  Syst.  Nat,  ed.  x,  1758;  ed.  xn,  p.  1098,  1766.  Gray 
(restr.,  Pars),  1840;  Synopsis,  p.  I,  1866.  Perrier  (Pars),  Revis.  Stell., 
op.  cit.,  iv,  p.  302,  1875.  Bell,  System.  Arrangement,  Proc.  Zool.  Soc. 
London,  1881,  p.  492. 

Stellonia  (pars)  NARDO,  Oken's  Isis,  p.  716,  1834.    Agassiz,  Prod.,  p.  191,  I835.1 

Vr -aster  (pars)  FORBES,  Mem.  Wernerian  Soc.,  vui,  p.  114,  1839. 

Asteracanthion  (pars)  MULLER  and  TROSCHEL,  Monatsb.  Akad.  wiss.,  Berlin, 
p.  102,  1840;  Syst.  Ast,  p.  14,  1842.  A.  Agassiz,  North  Amer.  Starfishes, 
p.  94,  1877  (structure). 

Asterias  NORMAN,  Ann.  and  Mag.  Nat.  Hist,  ser.  3,  xv,  p.  126,  1865. 

Asterias  (pars)  SLADEN,  Voy.  Chall.,  xxx,  p.  560,  1889.  Perrier,  Exp.  Trav. 
et  Talism.,  p.  108,  1894. 

Diplasterias  (Pars)  PERRIER,  Exp.  Trav.  et  Talism.,  p.  108,  1894. 

As  here  restricted,  this  genus  is  characterized  by  having  the  larger 
dorsal  ossicles  well  developed,  with  three  or  four  lobes,  united  in  a 
reticulated  or  irregular  manner,  and  with  oblong  transverse  ossicles, 
uniting  the  longitudinal  series,  and  thus  leaving  relatively  large, 
irregular  papular  areas,  from  which  arise  clusters  of  few  or  numer- 
ous small  papulae.  The  median  radial  dorsal  ossicles  usually  form  a 
distinct  row,  with  spines  more  prominent  than  the  others.  The 
inter actinal  plates  usually  consist  of  only  one  or  two  (sometimes 
three)  rows,  including  the  synactinal  or  connective  plates,  all  of 
which  are  usually  so  united  as  to  leave  papular  areas.  (See  p.  33 
above.) 

The  adambulacral  spines  may  stand  either  one  or  two  to  a  plate, 
but  very  commonly  they  are  alternately  or  irregularly  one  and  two. 
The  actinal  and  lower  marginal  spines  usually  form  three  to  five 
rows,  which  are  longer  and  larger  than  the  dorsal  spines,  and  in 
large  specimens  often  stand  two  or  three  on  one  plate.  The  supero- 
marginal  or  lateral  spines  generally  form  a  distinct  row,  leaving  a 
lateral  naked  lane  or  "  channel "  between  it  and  the  lower  marginals ; 
a  small  series  of  ossicles  may  sometimes  be  interpolated  in  this  area, 
at  the  bases  of  the  arms.  The  dorsal  spines  may  be  of  almost  any 
shape,  but  are  generally  shorter  than  the  marginals  and  actinals 
and  often  different  in  form.  The  spines  are  generally  surrounded 
by  groups  or  wreaths  of  forcipate  or  minor  pedicellariae,  but  similar 
pedicellariae  may  be  scattered  between  the  dorsal  spines,  or  among 
the  papulae,  or  they  may  form  dermal  clusters. 

*The  genus  Stellonia,  as  understood  by  L.  Agassiz  (Prod.,  p.  191,  1835), 
included  not  only  the  present  genus  Asterias  (sens,  ext),  but  also  Echinaster, 
Solaster,  Heliaster,  etc.  The  first  species  cited  was  S.  rttbens  (L.). 


IO2  VERRILL 

The  forficulate  or  major  pedicellariae  are  usually  lanceolate  or 
ovate,  and  more  or  less  compressed ;  they  may  be  scattered  between 
the  dorsal  spines,  around  the  papular  areas,  but  are  more  constantly 
present  on  the  ventral  interbrachial  areas,  and  on  the  naked  channel 
or  lane  between  the  upper  and  lower  marginal  spines ;  they  frequently 
occur  on  the  adambulacral  and  interactinal  spines,  where  they 
are  usually  of  smaller  size,  and  also  within  the  edge  of  the  ambu- 
lacral  groove,  attached  to  the  adambulacral  plates,  when  they 
usually  have  longer  pedicels.  They  generally  have  characteristic 
forms  in  each  species.  In  some  large  specimens  they  may  be  almost 
lacking,  though  abundant  in  others  of  the  same  species.  In  very 
young  specimens  they  are  usually  few  or  lacking. 

The  genital  ducts,  in  all  those  species  dissected,  are  connected  with 
a  pair  of  small  genital  pores  on  the  dorsal  side,  in  the  interbrachial 
areas. 

The  ambulacral  feet  and  pores  are  generally  in  four  rows,  but 
may  be  so  crowded  as  to  form  six  apparent  rows,  near  the  base  of  the 
rays.  These  forms  all  have  great  powers  of  restoration  and  repair 
after  injury,  being  able  to  replace  large  portions  or  all  of  the  disk 
as  well  as  part  or  all  of  the  rays,  but  spontaneous  fission  has  not 
been  observed  in  any  of  the  typical  species,  all  of  which,  so  far  as 
studied,  develop  from  a  free-swimming  brachiolarian  larva,  in  con- 
trast with  the  species  of  the  genus  Leptasterias,  which  have  no  free- 
swimming  stage,  for  their  eggs  and  larvae  are  attached  in  clusters  to 
the  oral  region,  till  capable  of  creeping.  They  have  ventral  genital 
pores. 

ASTERIAS  VICTORIANA  Verrill. 

Plate  Lin,  figure  I  (dorsal)  ;  plate  LJV,  figures  i,  2  (actinal  and  lateral)  ;  plate 
LXIX,  figure  4;  plate  LXXXII,  figures  i-ic  (details). 

Asterias  victoriana  VERMLL,  Amer.  Journ.  Sci.,  xxvin,  p.  68,  1909. 

Rays  five,  stout,  rather  rapidly  tapered.  Radii,  20  mm.  and 
95  mm.;  ratio,  1:475. 

Dorsal  skeleton  conspicuously  reticulated,  leaving  large  papular 
areas,  which  are  mostly  rounded  or  somewhat  elliptical,  the  trans- 
verse diameter  the  greater.  The  intervening  ossicles  are  strong  and 
prominent  above  the  surface,  as  narrow  convex  ridges ;  those  at  the 
intersections  and  in  the  radial  rows  larger  and  deeply  four-  to  six- 
lobed,  convex  in  the  middle,  with  a  central  mammilla  and  pit  where 
the  spine  is  attached. 

The  ossicles  of  the  two  marginal  rows  and  next  two  actinals  are 
thick,  nearly  equal  in  size  and  form,  and  proximally  stand  in  four 


SHALLOW-WATER  STARFISHES  I<>3 

or  five  regular  rows ;  the  upper  marginals  are  a  little  more  removed, 
but  the  others  are  closely  united  in  a  tesselated  manner,  leaving  only 
small  papular  pores  between  them.  The  exposed  part  is  convex, 
with  facets  and  pits  for  the  spines.  They  are  slightly  four-lobed, 
but  are  so  imbricated  that  they  appear  squarish  with  rounded 
corners,  or  ovate-triangular. 

The  dorsal  spines  consist  of  two  very  unequal  kinds.  The  larger 
ones  are  few  in  number  and  are  widely  scattered,  except  in  the 
median  radial  line,  where  they  form  a  pretty  regular  row ;  the  others 
stand  somewhat  in  quincunx,  but  may  belong  to  about  three  imper- 
fect rows  on  each  side.  These  spines  stand  on  the  larger  plates  at 
the  intersections  of  the  reticulations.  They  are  rather  large,  short, 
and  thick,  not  much  higher  than  broad,  with  enlarged,  truncate  or 
capitate  tips,  striated  on  the  sides  and  rough  on  the  top.  They  are 
about  1.5  mm.  broad.  Between  these  there  are  many  very  small 
inconspicuous  spines,  arranged  mostly  in  single  rows  along  the 
narrow  ossicles  that  form  the  sides  of  the  reticulations.  Some  of 
them  are  acute,  but  most  are  slightly  clavate  with  rough  or  spinulose 
tips.  Both  kinds  are  scattered  irregularly  on  the  central  area  of  the 
disk. 

Small  minor  pedicellariae  are  thickly  scattered  over  the  whole 
surface  between  the  spines  and  on  the  papular  areas,  and  also  form 
wreaths  around  the  larger  spines. 

The  superomarginal  spines  form  simple  regular  rows,  and  are 
much  like  the  large  dorsals  in  length  and  form,  but  are  smaller.  The 
intermarginal  channel  is  well  defined  and  of  moderate  width.  The 
inferomarginal  spines  form  a  regular  row,  mostly  simple,  but  fre- 
quently stand  two  on  a  plate  distally.  They  are  followed,  proxi- 
mally,  by  two  pretty  regular  close  parallel  rows  of  actinal  spines,  of 
about  the  same  size  and  shape.  These  three  rows  of  ventral  spines 
are  longer  than  the  superomarginals  and  less  clavate,  but  about  as 
stout.  They  are  blunt  and  sulcate  at  the  tips.  The  first  subactinal 
row  extends  only  to  about  the  end  of  the  proximal  third  of  the  ray; 
on  the  proximal  fourth  there  is  also  a  simple,  short  row  of  synactinal 
spines. 

The  synactinal  ossicles  are  small,  with  an  oblong  or  elliptical 
surface,  and  mostly  bear  a  single  spine ;  they  extend  only  to  about 
the  proximal  third  of  the  rays. 

The  adambulacral  spines  stand  two  on  a  plate,  or  else  in  certain 
parts  one  and  two  alternately,  thus  forming  two  or  three  crowded 
rows.  They  are  unequal,  not  very  slender,  the  inner  ones  slightly 
tapered,  the  outer  ones  stouter,  blunt,  as  long  as  the  ventral  spines, 


IO4  VERRILL 

but  more  slender.  They  increase  somewhat  in  length  and  thickness 
toward  the  mouth. 

The  two  apical  peroral  spines  are  rather  stouter  and  shorter  than 
the  adorals ;  their  side  spines  are  about  half  as  long  and  more  slender. 
The  epioral  spines  are  like  the  adorals. 

The  adoral  carina  is  rather  thick  and  stout,  composed  of  three 
pairs  of  contingent  plates  beyond  the  epiorals,  the  third  pair  bearing 
two  spines. 

Major  pedicellariae  of  moderate  size  occur  among  the  ventral 
spines  and  on  the  lateral  and  dorsal  surfaces,  but  are  not  numerous. 
They  are  compressed,  lanceolate  or  acute-ovate,  with  sharp  tips. 

The  type  of  this  species  was  from  near  Victoria,  B.  C.  (Prov.  Mus. 
B.  C.,  coll.  Dr.  C.  F.  Newcombe). 

ASTERIAS  POLYTHELA  Verrill. 

Plate  LV,  figures  i,  2  (dorsal)  ;  plate  LXX,  figure  8;  plate  LXXII,  figure  2;  plate 
LXXXIV,  figures  3,  4  (details)  ;  plate  LXXIX,  figures  1-20  (details). 

Asterias  polythela  VERRILL,  Amer.  Journ.  Science,  XXVIH,  p.  68,  1909. 

Rays  six,  stout,  of  moderate  length,  rounded  and  with  a  firm 
skeleton.  Radii,  20  mm.  and  80  mm. ;  ratio,  i :  4. 

Dorsal  surface  appears  rough  and  rugged.  It  bears  an  irregular 
number  of  large,  stout,  round  spines,  arranged  without  order,  except 
that  in  a  few  places  two  or  three  may  stand  in  a  median  series ;  else- 
where they  may  be  grouped,  two  to  five,  near  together,  or  stand 
singly.  These  spines  stand  on  raised  central  bosses  of  the  plates; 
they  are  constricted  somewhat  at  base  and  then  abruptly  enlarged 
below  the  middle ;  the  terminal  part  is  regularly  tapered  or  somewhat 
acorn-shaped  or  nipple-shaped,  longitudinally  finely  grooved,  ending 
in  a  blunt  apex.  They  are  2  mm.  to  4  mm.  high  and  1.5  mm.  to 
2  mm.  in  diameter.  Scattered  over  the  whole  surface  are  many 
small,  unequal,  short,  acorn-shaped  and  capitate  spines,  mostly  from 
2  mm.  to  4  mm.  in  diameter.  The  large  and  small  spines  are  all 
surrounded  by  close  wreaths  of  small  minor  pedicellariae ;  clusters  of 
these  are  also  attached  to  the  skin,  so  that  the  surface  appears  to  be 
almost  covered  with  them. 

The  marginal  and  actinal  rows  of  spines  are  pretty  regular  and 
smaller  than  the  dorsals.  The  upper  marginals  stand  mostly  one  to  a 
plate  proximally  and  two  to  a  plate  distally.  They  are  shaped  some- 
what like  the  large  dorsals  and  nearly  as  long,  but  only  about  half  as 
thick.  The  lower  marginals  are  about  as  long,  but  stouter;  they 


SHALLOW- WATER   STARFISHES  1 05 

stand  either  one  or  two  to  a  plate.  A  short  row  of  smaller  spines  is 
interpolated  between  the  upper  and  lower  marginals  proximally. 
The  peractinal  spines  are  like  the  lower  marginals  proximally  and 
form  a  regular  row,  one  to  a  plate.  The  adambulacral  spines  are 
small,  round,  blunt,  mostly  two  to  a  plate,  sometimes  one  in  certain 
parts,  divergent  and  almost  concealed  by  large  clusters  of  small, 
ovate,  major  pedicellariae  on  the  inner  ones,  and  clusters  of  major 
pedicellariae  on  the  outer  ones;  many  large  clusters  of  major  pedi- 
cellariae are  attached  to  the  inner  edge  of  the  plates  within  the  fur- 
row. A  few  much  larger,  blunt-ovate,  major  pedicellariae  with 
finely  denticulate  jaws,  occur  on  the  interradial  spaces  and  between 
the  proximal  marginal  spines. 

The  type  was  taken  off  the  Arctic  coast  of  Alaska  by  the 
U.  S.  R.  S.  "  Corwin  "  in  1885,  No.  16889  (U.  S.  Nat.  Mus.,  No. 
15820). 

ASTERIAS  NANIMENSIS  Verrill,  sp.  nov. 
Plate  LXI,  figures  i-ib  (dorsal  and  actinal  sides). 

Rays  five,  long,  narrow,  tapered,  nearly  half-round,  the  height 
about  equal  to  the  breadth  as  dried.  Radii,  180  mm.  and  18  mm.; 
ratios,  1 : 10.  Whole  upper  surface  of  rays  and  disk  pretty  regu- 
larly reticulated  by  the  lobed  plates  and  ossicles,  each  plate  bearing 
a  single  central  terete  spine,  1.5  mm.  to  2  mm.  high,  with  a  blunt 
fluted  tip.  These  spines,  on  the  rays,  are  arranged  nearly  in 
quincunx,  but  the  median  dorsal  row  is  distinct,  with  the  spines 
pretty  regular  and  close  together.  The  upper  marginal  row  is  dis- 
tinct but  scarcely  different  from  the  dorsals.  Between  the  upper 
marginals  and  the  median  row  there  might  be  reckoned  four  or  five 
zigzag  and  irregular  rows,  but  the  arrangement  is  rather  reticulate. 
The  dorsal  spine-bearing  plates  are  small,  with  a  central  boss  and 
usually  four  or  five  lobes,  giving  a  stellate  form,  the  lobes  united 
to  those  of  adjacent  plates  by  one  or  two  small  ossicles,  thus  leaving 
rather  large,  quadrangular,  rhombic,  or  pentagonal  papular  areas 
between  them.  Large  dense  clusters  of  minor  pedicellariae  surround 
the  bases  of  all  the  dorsal  and  marginal  spines,  and  the  outer  sides 
of  the  peractinal  spines.  /The  upper  marginal  plates  are  much  like 
those  above,  but  rather  stouter  and  in  a  more  regular  row;  their 
inferior  lobe  is  larger  and  stouter  than  the  others,  is  shoe-shaped, 
and  joins  the  upper  lobe  of  the  lower  marginal  plates,  usually  without 
an  intervening  ossicle,  leaving  a  well  defined  narrow  marginal  chan- 
nel  with  large  quadrangular  papular  areas  between  them.  The 


106  VERRILL 

lower  marginal  plates  are  similar  and  equal  in  number,  but  more 
regular.  Each  bears  a  single  spine  much  like  the  dorsals  in  size  and 
form.  Below  this  row  there  is  a  wide  naked  channel  with  large 
papular  areas,  like  those  above  it.  Between  this  row  and  the  adambu- 
lacral  plates  there  is  only  one  row  of  plates.  These  are  reckoned  as 
peractinal  plates.  They  correspond  in  number  and  size  with  the 
marginals  and  are  placed  close  to  the  adambulacrals,  which  they  join 
generally  with  no  connecting  ossicles  between;  when  rarely  such 
ossicles  occur,  they  are  few  and  very  small  or  rudimentary.  In  the 
narrow  channel  between  the  peractinal  and  adambulacral  plates 
there  is  a  row  of  small  papular  areas,  each  with  one  or  few  rounded 
papulae.  Each  peractinal  plate  bears  two  strong  equal  spines, 
decidedly  larger  and  longer  than  the  marginals,  the  proximal  ones 
2  mm.  to  3  mm.  long.  Their  tips  are  usually  flattened,  and  often 
enlarged  and  fluted  or  grooved. 

The  adambulacral  spines  are  numerous  and  regular,  two  to  a  plate, 
forming  two  close  rows ;  they  are  tapered,  blunt  or  subacute,  about 
four  pairs  corresponding  to  one  peractinal  plate.  Major  pedicellariae 
are  few  on  the  type.  Those  found  were  on  the  adambulacral  spines 
and  inner  margins  of  their  plates.  They  are  rather  large,  lanceolate, 
acute,  about  as  thick  as  the  adjacent  spines.  None  were  found  on 
the  back. 

The  disk  is  reticulated  much  like  the  rays,  with  large  angular 
interspaces.  The  madreporite  is  large  (5  mm.),  prominent,  with 
radiating  and  dichotomous  gyri.  It  is  surrounded,  in  the  type,  by 
seven  spines,  like  the  other  adjacent  spines. 

The  central  disk-spine  rises  from  a  five-lobed  plate.  It  is  sur- 
rounded by  five  spines,  and  beyond  this  with  a  circle  of  ten  spines. 

The  type  is  from  Departure  Bay,  Nanaimo,  British  Columbia, 
twenty-five  fathoms,  mud  (Chas.  H.  Young,  Canadian  Geological 
Survey,  1909).  fllfc;$/ 

This  species  seems  to  be  closely  related  to  the  genus  Orthas- 
terias,  in  several  respects,  and  especially  to  O.  kcehleri,  by  reason  of 
the  rather  indefinite  or  sublongitudinal  arrangement  of  its  dorsal 
spines,  in  about  seven  obscure  rows.  It  differs  from  the  latter  in 
the  shorter,  stouter,  fluted  and  crowded  dorsal  spines ;  the  psominent 
median  row ;  the  double  peractinal  row  of  elongated  spines ;  etc. 


SHALLOW-WATER   STARFISHES  IO7 

ASTERIAS  ACERVATA  Stimpson. 
Plate  xxvii,  figures  1, 2  (var.  acervata,  type)  ;  plate  cvi,  figure  3  (var.). 

Asterias  rubens  (pars)  and  A.  minuta  (pars)  FABRICIUS,  Fauna  Groenl.,  pp. 
367,  370,  1780  (non  LiNNfc). 

Asterias  violacea  SABINE,  Suppl.  Parry's  Voyage,  p.  ccxxni,  1824  (non  Miiller). 

Asteracanthion  polaris  MULLER  and  TROSCHEL,  Syst.  Aster.,  p.  16, 1842  (young). 
Steenstrup,  1855,  p.  240.  Liitken,  Vidensk.  Meddel,  1857,  pp.  28,  29,  1857. 
Perrier,  Ann.  Sci.  Nat.,  Paris,  pp.  33-36,  pL  i,  fig.  6,  1869.  Duncan  and 
Sladen,  1878,  pp.  265,  266,  1878;  ditto,  Echinod.  Arctic  Sea,  pp.  23-27,  pi. 
n,  figs.  4-8,  1881. 

Asterias  polaris  STIMPSON,  Proc.  Boston  Soc.,  vm,  p.  271,  1862.  Verrill,  Proc. 
Boston  Soc.  Nat.  Hist,  x,  p.  356,  1866;  1867,  p.  268;  Amer.  Journ.  Sci., 
xi,  p.  420,  1876;  Amer.  Journ.  Science,  XLIX,  p.  208,  1895;  in  Packard, 
Fauna  of  Labrador,  p.  268,  1867.  Perrier,  Arch.  Zool.  Exper.,  pp.  332, 
357.  J875  (Greenland  examp.  described).  Bush,  Proc.  U.  S.  Nat.  Mus., 
vi,  p.  246,  1883.  Sladen,  Voy.  Challenger,  xxx,  p.  826,  1889.  Ludwig, 
Fauna  Arctica,  I,  p.  485,  1900  (distribution),  non  Sabine,  1824. 

Asterias  acervata  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vm,  p.  271,  1862. 
Bell,  op.  cit,  1881,  p.  494.  Murdoch,  Report  International  Polar  Exped., 
p.  158,  1885.  Ludwig,  Echinod.  des  Beringsmeeres  Zool.  Jahrb.,  p.  287, 
1886;  Fauna  Arctica,  i,  p.  485,  1900.  Sladen,  Voy.  Chall.,  xxx,  p.  818, 
1889.  Bell,  1881,  p.  494. 

'Asterias  borealis  PERRIER,  Arch.  Zool.  Exper.,  iv,  pp.  323,  357,  1875  (descr. 
Labrador  example).  Bell,  op.  cit.,  1881,  p.  497. 

Asterias  douglasi  PERRIER,  Revis.,  Arch.  Zool.  Experim.,  iv,  pp.  333,  357, 
1875  (based  in  part  on  one  of  Gray's  supposed  cotypes  of  A.  katherinee  = 
A.  polaris,  teste  Perrier,  in  note,  p.  357).  Bell,  Arrangement,  p.  497,  1881. 

Of  the  true  Atlantic  polaris1  I  have  examined  large  numbers  of 
examples,  in  various  collections,  and  have  also  collected  it  myseli  on 
the  southern  coast  of  Labrador  and  at  Anticosti  Island  (1861), 
where  it  is  common  at  low  tide  among  the  rocks,  often  associated  in 
that  region  with  A.  vulgaris,  which  it  resembles  in  size  and  color. 

Among  hundreds  of  specimens,  I  have  never  seen  one  with  five 
rays,  though  such  are  said  to  occur  very  rarely. 

The  size,  form,  and  arrangement  of  the  dorsal  spines  vary  widely, 
as  in  all  the  allied  species,  but  still  the  general  appearance  is  pretty 
characteristic.  The  dorsal  spines  are  always  numerous,  capitate  (or 
when  more  slender,  clavate),  and  always  more  or  less  clustered  or 
acervate,  unless  in  very  young  examples. 

In  large  Labrador  specimens,  seven  to  nine  inches  in  diameter 
(larger  radii  may  be  100  mm.  to  150  mm.),  the  dorsal  spines 

*This  form  should  be  called  Asterias  acervata  borealis,  since  A.  polaris 
is  a  preoccupied  name. 


IO8  VERRILL 

become  much  more  numerous  and  more  conspicuously  clustered 
than  in  those  of  the  ordinary  sizes. 

The  upper  and  lower  marginal  spines  are  conspicuously  longer 
than  the  dorsals  and  are  usually  tapered.  They  commonly  stand 
singly  on  each  plate  and  form  a  simple  row  in  each  series  on  the 
basal  part  of  the  rays,  but  distally  the  upper  row  usually  becomes 
double,  with  two  spines  on  each  plate.  A  short  lateral  row  is  often 
interpolated  between  the  two  marginal  series. 

The  actinals  are  like  the  marginals  and  form  one  continuous  and 
usually  simple  row,  close  to  the  lower  marginals.  The  marginal  and 
actinal  plates  are  stout,  wide,  and  closely  joined. 

The  subactinal  spines  are  rather  smaller  and  do  not  usually  extend 
beyond  the  basal  third  of  the  rays ;  they  are  close  to  the  adambula- 
crals.  The  latter  are  biserial,  but  an  occasional  plate  bears  but  one 
spine;  they  are  slender,  tapered,  and  terete,  and  carry  conspicuous 
clusters  of  minor  pedicellariae. 

The  major  pedicellariae  of  the  Labrador  borealis  are  very  variable 
in  size,  just  as  Stimpson  describes  those  of  acervata.  Those  scat- 
tered on  the  back  are  very  small,  short-ovate;  those  on  the  lateral 
channels  are  much  larger,  but  not  unusually  large,  and  vary  from 
short-ovate  to  triangular-ovate ;  most  of  them  are  short,  with  very 
obtuse  tips,  and  are  not  distinctly  serrate  externally.  Some  speci- 
mens have  only  a  few  of  these  pedicellariae,  and  of  smaller  size  than 
usual. 

The  color  in  life  varies  from  pale  red  or  pink  to  purple  and  pale 
violet. 

The  name  A.  polaris  is  preoccupied  by  Asterias  polaris  Sabine, 
1824. 

The  original  type  of  Dr.  Stimpson's  Asterias  acervata  is  still  pre- 
served in  the  U.  S.  National  Museum.  Photographs  of  it  have  been 
sent  to  me  by  Dr.  R.  Rathbun  for  reproduction.  (See  pi.  xxvn, 
figs,  i,  2.) 

Dr.  Stimpson  gave  the  following  description : 
"  Rays  six,  more  convex  and  more  tapering  than  in  A.  polaris. 
Disc  of  moderate  size.  Proportion  of  the  diameters,  1 : 4.5.  Ambu- 
lacral  spines  in  two  rows  (two  to  each  plate),  rather  stout,  cylin- 
drical, and  thickly  covered  with  minor  pedicellariae  near  the  tips. 
Ventral  spines  standing  in  two  or  three  irregular  rows.  Lateral 
channel  not  well  marked,  and  sometimes  occupied  by  very  small 
spines.  Lateral  spines  standing  singly  in  one  row,  and  more  pointed 
than  the  ventrals.  Dorsal  spines  more  numerous  and  crowded  than 


SHALLOW-WATER   STARFISHES  ICX) 

in  A.  polaris,  and  of  greater  diversity  in  size,  the  larger  ones  being 
collected  in  heaps  which  form  three  regular  longitudinal  rows  on  the 
rays.  These  large  spines  are  capitate,  with  obtusely  conical  and 
striated  heads;  there  are  usually  about  six  spines  in  each  heap,  the 
central  one  being  much  the  largest  and  overtopping  the  others,  which 
form  a  circle  around  it.  The  small  spines  between  the  heaps  are 
quite  uniform  in  size,  and  have  globular  tips.  Disc  surrounded  by  a 
ring  of  six  heaps  of  spines,  within  which  there  is  sometimes  another 
ring  of  the  same  number,  and  always  a  heap  in  the  middle.  Madre- 
poric  plate  surrounded  with  a  circular  canal  and  a  ring  of  thirteen 
spines.  All  the  spines,  both  ventrals  and  dorsals,  are  surrounded  by 
minor  pedicellariae,  as  in  A.  polaris.  The  major  pedicellariae,  which 
are  most  numerous  on  the  sides  of  the  rays,  are  scattered,  and  very 
irregular  in  size,  varying  from  one  two-hundredth  to  one-twentieth 
of  an  inch  in  length;  the  largest  ones  are  stout,  as  long  as  broad, 
conical,  or  almost  globular,  having  valves  with  broad,  dentated 
extremities.  Papulae  numerous,  scattered,  and  often  forming  groups. 
Color,  in  life :  above,  clouded  with  very  dark  brown ;  madreporic  plate 
cream-colored.  Sides  of  rays,  and  inferior  surface,  of  a  yellowish 
cream-color.  Diameter,  five  and  a  half  inches. 

"  Habitat,  Behring's  Straits,  on  clean  gravelly  bottoms,  in  from 
five  to  fifteen  fathoms.  U.  S.  North  Pacific  Expedition.  Wm. 
Stimpson." 

Dr.  Stimpson  was  the  naturalist  of  the  North  Pacific  Exploring 
Expedition,  under  Ringold  and  Rogers,  and  therefore  probably  made 
his  notes  on  the  colors  from  personal  observation  of  living  specimens. 
I  have  received  from  the  U.  S.  National  Museum  a  specimen  from 
Cape  Smith,  Alaska  (No.  7630),  which  agrees  very  closely  with 
Stimpson's  type.  The  radii  are  15  mm.  and  55  mm.  The  dorsal 
spines  are  formed  as  described  by  Stimpson  and  are  conspicuously 
acervate ;  but  though  the  clusters  are  somewhat  in  three  radial  rows 
on  the  rays,  the  rows  are  decidedly  irregular.  Between  the  clusters 
the  ossicles  are  obviously  areolate  or  reticulate,  and  bear  small  capi- 
tate spines.  The  upper  marginal  spines  are  about  equal  in  thickness 
to  the  larger  dorsals  and  rather  longer ;  they  stand  either  one  or  two 
on  a  plate.  The  lower  marginals  are  longer  but  not  so  stout,  and 
form  a  regular  row,  one  to  a  plate.  There  is  a  short  intermarginal 
row  of  smaller  spines  proximally.  The  peractinals  are  similar  to  the 
lower  marginals  and  form  a  regular,  simple  row  close  to  the  latter. 
Subactinal  row  short.  The  adambulacral  spines  are  irregularly 
diplacanthid ;  they  are  moderately  long,  scarcely  tapered,  obtuse. 


IIO  VERRILL 

Cape  Smith,  Alaska  (Murdoch,  Point  Barrow  Expedition,  1884). 

A  young  specimen  of  acervata  (U.  S.  Nat.  Mus.,  No.  6082),  from 
Nazan  Bay,  Alaska,  collected  by  Dr.  W.  H.  Dall,  in  1872,  at  low 
tide,  differs  considerably  from  the  type.  This  is  probably  due  largely 
to  immaturity.  The  radii  are  16  mm.  and  48  mm.  The  principal 
dorsal  spines  are  short,  capitate,  and  form  numerous  small,  but  con- 
spicuous raised  clusters  on  the  disk  and  median  zone  of  the  rays,  but 
are  lacking  on  the  superolateral  surfaces,  where  there  are  two  pretty 
regular  rows  of  very  small  capitate  spines  and  some  scattered.  The 
median  clusters  are  so  crowded  that  they  are  often  in  contact  proxi- 
mally.  The  two  marginal  and  the  peractinal  rows  are  regular,  one 
to  a  plate,  except  that  the  upper  marginals  are  double  distally.  The 
upper  marginals  are  short,  capitate,  nearly  as  large  as  the  larger  dor- 
sals ;  those  of  the  lower  marginal  and  peractinal  rows  are  larger,  not 
so  stout,  clavate  or  subclavate.  The  adambulacral  spines  are  numer- 
ous, subdiplacanthid ;  distally  they  are  clavate,  but  become  longer, 
more  slender,  and  subacute  near  the  mouth.  The  pedicellariae  are 
similar  to  those  of  the  preceding  form,  but  less  numerous ;  the  large 
dermal  ovate-lanceolate  pedicellariae  are  present.  The  apical  plate  at 
the  tips  of  the  rays  is  rather  large  and  the  small  spines  are  thickly 
clustered  around  it. 

Ludwig  (op.  cit,  1900)  suggested  that  acervata  is  identical  with 
A.  camtschatica  Brandt  (op.  cit.,  1835).  If  this  could  be  proved,  the 
latter  name  would  have  priority  over  all  the  others.  But  Brandt's 
description,  which  was  based  on  a  colored  drawing  of  a  young  or 
small  six-rayed  starfish  (diameter  of  disk  one  inch,  length  of  rays 
one  to  one  and  one-fourth  inches),  is  far  too  imperfect  to  be  of 
much  value  by  itself  for  the  identification  of  a  species  of  this  genus. 

I  have  elsewhere  given  as  good  or  better  reasons  for  identifying 
his  species  rather  with  a  different  one  (A.  multiclava)  found  on  the 
Siberian  coast  (see  p.  114),  but  his  description  would  apply  as  well, 
also,  to  forms  of  L.  epichlora. 

On  the  Atlantic  coast  this  Arctic  species  extends  southward  to  the 
Gulf  of  St.  Lawrence,  at  Anticosti  Island,  etc.,  and  to  Newfoundland 
and  the  fishing  banks  off  Nova  Scotia,  but  not  to  New  England. 
It  is  common  on  the  Labrador  and  Greenland  coasts  as  far  north  as 
north  latitude  70°,  at  least.  On  the  Pacific  side  it  is  found  in  the 
Arctic  Ocean  and  on  the  coasts  of  Bering  Sea  and  the  Aleutian 
Islands,  but  so  far  as  positively  known  to  me  it  does  not  occur  on  the 
southwest  coast  of  Alaska.  It  is  also  known  from  Siberia.  A 
typical  specimen  of  acervata,  received  from  the  U.  S.  National 


SHALLOW- WATER   STARFISHES  III 

Museum  (No.  7630),  was  found  on  the  beach  at  Cape  Smith, 
Alaska,  October,  1881,  by  Mr.  Murdoch  (Point  Barrow  Expedi- 
tion). The  young  specimen  described  above  was  from  Nazan, 
Alaska  (W.  H.  Dall). 

Ludwig  (op.  cit.,  1886,  p.  287)  refers  to  A.  acervata  as  follows: 

"  Of  this  species,  which  has  become  known  through  Stimpson, 
there  are  a  number  of  examples,  the  largest  of  which  measures 
17.5  mm.  The  specimens  come  from  Bering  Strait,  from  a  depth  of 
from  five  to  fifteen  fathoms,  and  according  to  their  exact  localities 
are  divided  as  follows :  Two  specimens  from  Lorenz  Bay ;  one  full- 
grown  and  one  quite  young  specimen  from  Emma  Harbor,  Plover 
Bay;  one  specimen  from  St.  Matthew's  Island;  one  specimen  from 
northwest  of  St.  Matthew's  Island,  at  a  depth  of  twenty-five 
fathoms ;  one  specimen  from  St.  Paul's  Island ;  three  specimens  with- 
out definite  locality,  from  a  depth  of  from  twenty-three  to  twenty- 
five  fathoms.  All  the  foregoing  specimens  are  six-armed;  and  fur- 
ther, one  five-armed  specimen  from  St.  Paul's  Island." 

It  seems  probable  that  Stimpson's  A.  acervata  is  not  specifically 
distinct  from  the  North  Atlantic  and  Arctic  A.  polaris.  I  have 
compared  typical  specimens  of  it  with  specimens  of  A.  polaris  from 
Labrador  and  found  a  close  agreement,  so  that  there  can  scarcely  be 
a  doubt  of  their  identity,  though  they  may  be  separated  as  geographi- 
cal varieties.  In  that  case  the  North  Atlantic  form  should  be  called 
'Asterias  acervata  borealis  (Per.). 

Stimpson's  description  applies  fairly  well  to  some  specimens  of 
the  same  size  from  the  Atlantic,  except  in  one  particular.  He 
mentions  the  occasional  occurrence  of  stout,  serrate  major  pedicel- 
lariae  in  the  lateral  channels,  evidently  referring  to  the  form  char- 
acteristic of  the  various  species  of  Pisaster,  and  also  found  in 
Leptasterias  epichlora  alaskensis  Ver.  and  var.  subnodulosa  Ver.  I 
have  not  been  able  to  find  such  pedicellariae  on  the  Atlantic  A.  polaris. 
Therefore,  it  is  not  unlikely  that  some  of  Stimpson's  specimens 
belonged  to  subnodulosa,  which  is  also  six-rayed  and  acervate  and 
often  has  much  resemblance  to  his  A.  acervata.  Moreover,  it  is  a 
very  common  littoral  variety  on  the  Alaskan  coast,  and  Stimpson 
could  hardly  have  failed  to  have  had  it  in  his  possession.  But  in 
other  respects  his  description  certainly  does  not  apply  to  the  latter. 


112  VERRILL 

ASTERIAS  KATHERINJE  Gray  (non  Perrier). 

Plate  LI,  figures  i,  2;  plate  LII,  figure  i  (dorsal,  large)  ;  plate  LXXXIII,  figure  I 

(details). 

Asterias  katherina  GRAY,  Annals  and  Mag.  Nat.  Hist,  vi,  p.  179,  1840;  Synop- 
sis, Genera  and  Species,  p.  2,  1866  (non  Perrier,  1875). 

Asteracanthion  katherina  MULLER  and  TROSCHEL,  Syst  Aster.,  p.  19,  1843 
(translation  from  Gray).  Dujardin  et  Hupe,  Echinod.,  in  Suites  a 
Buffon,  p.  339,  1862  (translation). 

Gray's  brief  description  of  A.  katherina  is  as  follows: 

"  Rays  six  or  rarely  five,  nearly  three  times  as  long  as  the  width 
of  the  body;  back  with  scattered  and  crowded,  blunt,  rough-tipped 
spines."  He  placed  it  in  his  section  having  the  adambulacral  spines 
crowded  "  as  if  two-  or  three-rowed,"  with  the  back  "  netted," 
ventral  spines  in  two  or  three  rows;  lateral  spines  in  a  single  row. 
He  gave  the  locality  as  "  mouth  of  the  Columbia  River,  Lady 
Katherine  Douglas." 

To  this  species,  long  misunderstood  and  little  known,  I  refer  with 
confidence  a  good  specimen  obtained  by  Mr.  A.  Agassiz  in  the  Gulf 
of  Georgia,  about  1860  (No.  1181,  Mus.  Comp.  Zool.  See  plates 
LI,  LII.)  It  agrees  in  all  respects  with  Gray's  brief  description. 

Rays  six,  moderately  stout,  evenly  convex  above  and  regularly 
tapered ;  larger  radius,  85  mm. ;  shorter,  18  mm. ;  ratio,  i :  4.72. 

The  dorsal  spines  are  very  numerous,  mostly  rather  small,  none 
large.  Smaller  ones  clavate;  larger  ones  somewhat  capitate; 
arranged  crowdedly  without  any  regular  order,  but  reticulate  or  in 
short  rows  in  many  places,  both  on  the  disk  and  rays,  not  acervate 
and  not  conspicuously  diverse  in  size,  though  many  on  the  transverse 
ossicles  are  quite  small.  The  median  dorsals  are  like  the  rest  and 
do  not  form  an  evident  row. 

The  superomarginals  are  a  little  longer  and  more  cylindric,  obtuse 
or  slightly  clavate,  placed  in  a  regular  row,  one  to  each  plate. 

The  inferomarginals  are  quite  similar  and  stand  mostly  one  to  a 
plate,  but  occasionally  there  are  two  to  a  plate. 

The  interactinal  spines  are  shorter  and  stouter  than  the  marginals, 
cylindric  or  slightly  clavate,  obtuse  or  pinched  at  the  tip,  nearly 
smooth.  They  form  two  close  rows,  or  three  in  some  places,  proxi- 
mally,  crowded  between  the  inferomarginals  and  adambulacrals, 
with  very  small  papular  areas  around  and  between  them. 

They  apparently  stand  on  two  close  rows  of  interactinal  plates, 
for  there  are  small  papular  pores  between  the  rows.  The  subactinal 
row  extends  in  most  cases  only  about  half  the  length  of  the  ray. 


SHALLOW- WATER   STARFISHES  113 

The  adambulacral  spines  are  unusually  numerous,  crowded  in 
three  or  more  rows.  They  stand  mostly  two  on  a  plate,  but  often 
there  are  three  on  part  of  the  plates,  or  alternately  two  and  three. 
They  are  of  moderate  length,  the  outer  ones  longer,  terete,  slightly 
clavate,  obtuse;  the  inner  ones  shorter,  acute.  Oral  spines  stouter, 
a  little  curved. 

The  minor  pedicellariae  are  very  small  and  form  small  wreaths  on 
the  dorsal  and  superomarginal  spines.  The  major  pedicellariae  are 
very  numerous  and  small,  ovate,  those  on  the  dorsal  side  are  unus- 
ually small  and  thickly  scattered  over  the  dermal  areas.  On  the 
ventral  spines  they  are  little  larger  and  more  acute,  numerous  on  the 
spaces  between  the  actinal  and  marginal  and  adambulacral  spines, 
and  also  form  clusters  on  the  spines.  Those  on  the  lateral  areas  are 
rather  larger  and  more  lanceolate,  but  still  smaller  than  in  most 
species.  All  the  papular  areas  are  small.  The  madreporic  plate  is 
large,  with  very  numerous  gyri. 

Gulf  of  Georgia  (A.  Agassiz,  1860,  Mus.  Comp.  Zool.,  No.  1181). 

This  species,  as  here  described,  resembles  A.  borealis  in  form,  but 
it  has  smaller  and  more  numerous  dorsal  spines,  not  acervate,  and 
the  actinal  and  adambulacral  spines  are  more  numerous  and  more 
crowded.  The  pedicellariae  are  also  different. 

M.  Perrier  (1875,  p.  332)  stated  that  he  found  in  the  British 
Museum  a  number  of  specimens,  some  of  which  probably  were 
Gray's  types  (though  the  label  was  loose).  Most  of  these  were  six- 
rayed  and  monacanthid,  but  two  were  five-rayed.  With  them  was 
also  a  single  specimen  of  a  six-rayed  diplacanthid  species  that 
Perrier  referred  to  A.  douglasi,  which  last  Bell,  Perrier  (1881),  and 
Ludwig  ( 1900)  referred  to  A.  polaris  —  borealis. 

It  is  evident  that  Gray's  name  must  be  applied  to  a  diplacanthid 
species,  not  to  the  monacanthid  species  to  which  Perrier  restricted  it, 
and  which  I  have  named  A.  grayi.  If  it  were  really  certain  that  the 
specimens  found  by  Perrier  were  Gray's  cotypes,  the  name  should 
be  applied  either  to  the  six-rayed  one  called  douglasi  by  Perrier,  or 
to  the  five-rayed  specimens  referred  to  by  him.  But  as  the  label  was 
loose,  it  may  well  have  been  misplaced  in  the  long  interval  of  time 
(thirty-five  years),  or  other  specimens  may  have  been  added  to 
the  original  lot.  Such  accidents  happen  in  all  museums.  It  is  not 
unlikely  that  the  douglasi  Per.  of  the  British  Museum  lot  is  the  same 
as  the  species  described  by  me  as  katherina  above. 


1 14  VERRILL 

If  A.  douglasi*  be  really  identical  with  A.  borealis,  as  some  claim, 
and  that  particular  specimen,  seen,  but  not  described,  by  Perrier,  is 
the  same,  it  could  hardly  have  been  from  the  mouth  of  the  Columbia 
River,  as  stated  by  Gray;  for  A.  borealis  (or  acervata)  has  not  been 
found  on  the  northwest  coast  south  of  the  Aleutian  Islands  by  any  of 
the  numerous  recent  expeditions  to  that  coast,  so  far  as  I  know.  On 
the  other  hand,  the  monacanthid,  six-rayed  species  (grayi  V.) 
belongs  to  a  group  of  allied  species  well  represented  in  Puget  Sound 
and  southward,  so  that  the  locality  given  by  Gray  might  be  right  for 
that  one.  What  his  five-rayed  specimens  were  I  do  not  know;  pos- 
sibly they  were  E.  troschelii. 

ASTERIAS  MULTICLAVA  Verrill,  sp.  nov. 

V 

Plate  LVIII,  figure  2;  plate  LIX,  figure  i  (large,  actinal)  ;  plate  LXIX,  figure  i; 
plate  LXXXIV,  figures  2,  20  (details). 

??  Asterias  camtschatica  BRANDT,  Prod.,  p.  270,  1835  (description  insufficient, 
perhaps  A.  epichlora,  six-rayed  var.).  ?  Stuxberg,  Evert.  Fauna  Sib.  Is., 
p.  28,  1880.  Sladen,  Voy.  Chall.,  p.  820,  1889.  Ludwig  (pars},  Fauna 
Arctica,  i,  p.  485,  1900  (distribution). 

t?  Asteracanthium  canttschaticum  BRANDT,  in  Middendorff,  Reise,  n,  p.  32, 
185.1.  ?  Grube,  Nova  Acta  Acad.  Cxs.  Leop.,  xxvii,  pp.  23,  26,  1857 
(A  steracanthion) . 

Rays  six,  rather  long  and  tapered ;  disk  small ;  radii,  14  mm.  and 
62  mm. ;  ratio,  i :  4.4.  Dorsal  and  ventral  spines  short,  clavate,  and 
capitate,  very  numerous,  arranged  largely  in  distinct  radial  bands. 
The  dorsal  spines  are  all  similar  in  form  and  not  very  diverse  in  size. 
They  form  three  or  five  crowded  rows,  with  some  scattered  between ; 
the  median  row  is  a  little  more  conspicuous  and  its  spines  are 
crowded  and  a  little  clustered  in  many  places.  Those  of  the  disk 
form  many  small  clusters,  but  are  not  acervate;  a  circle  of  the 
smaller  ones  surrounds  the  madreporite.  The  superomarginal  row 
is  regular  and  compressed,  having  either  two  or  (in  the  adult)  three 
capitate  spines  to  a  plate.  The  inferomarginals  are  longer,  stout, 
clavate,  and  stand  either  one  or  two  to  a  plate,  according  to  the  age. 
The  peractinals  are  similar,  and  stand  mostly  one  (in  the  young), 
but  often  two  (in  the  adult),  to  a  plate;  there  is  also  a  short  sub- 
actinal  row,  and  a  short  proximal,  intermarginal  row  of  small  spines. 
Thus  in  the  adult  the  lateral  and  interactinal  spines  are  unusually 

1  The  types  of  A.  douglasi  are,  apparently,  the  five  specimens,  without  locality, 
preserved  in  the  Paris  Museum.  The  British  Museum  specimen  may  not 
have  been  identical,  but  really  Gray's  type,  from  off  the  Columbia  River. 


SHALLOW-WATER   STARFISHES  11$ 

numerous  as  well  as  the  dorsals.  In  two  small  specimens,  with  the 
larger  radii  24  mm.  and  28  mm.,  the  inferomarginal  and  the  per- 
actinal  rows  are  single,  but  the  superomarginal  rows  are  double. 
The  spines  in  these  are  less  clavate.  The  adambulacrals  are  irregu- 
larly diplacanthid  ;  distally  they  are  slender  and  subclavate,  obtuse  ; 
proximally  they  become  much  more  slender,  longer,  tapered,  sub- 
acute.  They  have  large  clusters  of  minor  pedicellariae,  with  some 
small  major  pedicellariae  intermixed.  Small  clusters  of  minor  pedi- 
cellariae are  also  found  on  the  lateral  and  dorsal  spines.  Dermal 
major  pedicellariae  of  small  and  moderate  sizes  are  numerous  between 
the  lateral  rows  of  spines.  They  are  ovate  and  ovate-lanceolate  in 
form,  and  subacute;  others,  much  smaller  and  mostly  ovate,  are 
found  with  them  and  also  on  the  dorsal  surface.  The  papular  areas 
are  small.  Color  in  life  :  "  Above,  deep  olive  green  with  the  tips  of 
the  spines  whitish;  those  of  the  edges  very  pink"  (Stejneger). 
The  type  (No.  15841,  U.  S.  Nat.  Mus.)  was  taken  February  2, 
1883,  at  Bering  Island,  Commander  Is.,  by  Dr.  L.  Stejneger.  Three 
smaller  specimens  are  from  Petropaulski,  Siberia,  Afbatross  col- 
lection. 

This  species  is  evidently  related  to  A.  acervata  borealis.  V  It  differs 
mainly  in  the  pretty  regular  arrangement  of  the  dorsal  spines  in  three 
or  five  multiple  rows  ;  in  having  the  superomarginal  rows  of  plates 
double-  or  triple-spined,  and  the  inferomarginals  and  peractinals 
double-spined  when  adult.  These  features  I  have  not  seen  in  the 
Atlantic  borealis,  nor  in  the  Pacific  form,  A.  acervata.  The  young 
specimens  resemble  some  species  of  Leptasterias,  like  L.  dispar. 

The  species  very  imperfectly  described  by  Brandt,  from  Siberia, 
has  always  been  doubtful,  Ludwig  considered  it  the  same  as 
A.  acervata  Stimpson  (see  p.  no),  but  that  does  not  seem  very  prob- 
able. It  is  more  likely  to  have  been  this  species,  if  either. 

Brandt's  original  description  was  based  on  a  colored  drawing  of 
a  very  young  specimen  ("  diameter  of  disk,  one  inch;  length  of  rays, 
one  to  one  and  one-fourth  inches  "). 

Aside  from  the  size  and  color,  the  only  other  characters  given  are 
that  the  dorsal  spines  ("papillae")  are  numerous  and  "truncate- 
capitate,"  reticulate,  and  form  "  subangular"  series  "  on  the  rays. 
The  last  statements  do  not  apply  very  well  to  A.  acervata,  but  would 
apply  very  well  to  this  species,  and  also  to  some  of  the  six-rayed 
varieties  of  epichlora.  But  the  latter  is  not  known  to  me  from  the 
Arctic  Ocean  nor  from  Siberia, 

The  capitate  form  of  the  larger  spines  is  characteristic  of  all  these 
forms,  as  well  as  many  others. 


7  .Juw&u**  q_  t  t^^Mt^  &< 


1 16  VERRILL 

On  the  whole,  especially  considering  the  locality,  it  is  more  prob- 
able that  Brandt's  original  type  of  A.  camtschatica  was  the  same  as 
the  present  species;  but  the  description  is  so  indefinite  and  vague 
that  it  seems  unsafe  to  use  his  name,  for  the  species  of  this  group 
are  very  numerous. 

Genus  Leptastcrias  Verrill. 

Leptasterias  VERBILL,  Proc.  Boston  Soc.  Nat.  Hist.,  x,  p.  350,  1866.  Type,  L. 
mulleri  Sars.  Sladen,  Voy.  Chall,  xxx,  p.  563,  1889.  Perrier,  Exp.  Trav. 
et  Talism.,  p.  108. 

An  extensive  group  of  small,  more  or  less  diplacanthid  starfishes, 
closely  related  to  typical  Asterias,  usually  with  a  single  row  of  inter- 
actinal  plates  and  spines.  Rays  five  or  six.  The  more  typical  forms 
have  slender  rays  and  a  small  disk,  with  the  dorsal  ossicles  irregularly 
arranged ;  others  have  several  subimbricated  radial  rows,  with  clus- 
tered spines.  It  differs  especially  from  typical  Asterias  in  the  dimin- 
ished number  and  larger  size  of  the  dorsal  and  lateral  papulae,  and  the 
small  number  of  interactinal  plates,  of  which  there  is  generally  only  a 
single  row.  The  eggs  and  young  are  carried  in  clusters,  adhering  in 
front  of  or  around  the  mouth  in  most  if  not  all  the  species.  The  genital 
pores  are  on  the  actinal  side,  near  the  mouth.  The  oviduct  is  wide 
and  short;  the  ovarian  tubules  are  few,  thick,  beaded  by  the  eggs. 
(See  also  p.  8).  Type,  L.  mulleri  (Sars). 

The  most  abundant  species  of  Alaska  (L.  epichlora)  is  by  no 
means  a  typical  member  of  this  group,  for  in  many  respects  it  is 
intermediate  between  it  and  typical  Asterias,  especially  in  having 
larger  clusters  of  papulae  and  often  two  proximal  rows,  or  sometimes 
three,  of  interactinal  spines  when  of  large  size. 

The  species  of  this  group  are  remarkably  variable  in  many  cases. 
This  is  particularly  true  of  those  inhabiting  the  vast  extent  and 
varied  localities  of  the  Northwest  coast. 

This  unusual  tendency  to  form  marked,  more  or  less  localized 
varieties  is,  like  the  same  phenomenon  seen  in  the  genus  Henricia, 
probably  due  to  the  fact  that  most,  if  not  all,  of  the  species  of  these 
genera  carry  their  young,  until  they  reach  the  starfish  form,  attached 
to  the  region  about  the  mouth,  and  consequently  have  no  free- 
swimming  larval  stages  (brachiolaria) ,  such  as  we  find  in  the 
species  of  typical  Asterias.  The  free-swimming  young  of  the  latter 
may  remain  afloat  many  days,  or  even  weeks,  and  thus  they  may  be 
carried  long  distances  by  the  currents,  in  one  generation,  or  they 
may  be  scattered  in  different  directions  over  large  areas,  thus  mixing 
up  the  young  from  various  localities. 


SHALLOW- WATER    STARFISHES  117 

On  the  contrary,  the  sessile  condition  of  the  young  in  the  species 
of  Leptasterias  insures  greater  safety,  and  though  the  eggs  are  much 
fewer  in  number,  a  far  greater  per  cent  will  reach  maturity. 

This  method  of  propagation  precludes  the  rapid  diffusion  of  such 
species,  for  they  can  only  migrate  by  means  of  the  slow  method  of 
creeping  by  the  use  of  their  ambulacral  feet,  unless  accidentally 
carried,  attached  to  floating  objects.  Under  these  conditions  any 
local  variations  that  may  arise,  unless  harmful,  are  likely  to  be  per- 
petuated by  inheritance  and  isolation.  As  a  matter  of  fact,  many  such 
varieties  are  already  known,  some  of  which  may  be  quite  localized, 
so  far  as  known,  while  others,  probably  of  much  earlier  origin,  are 
diffused  from  Puget  Sound  to  the  Aleutian  Islands. 

Very  extensive  collections  along  the  whole  coast  are  needed  before 
the  real  status  of  many  of  the  varieties  in  this  group,  as  well  as  in 
Henricia  and  Pteraster,  all  of  which  carry  their  young,  can  be  settled. 

Our  present  collections  are  quite  inadequate.  The  most  that  we 
can  now  do  is  to  describe  and  figure  those  varieties  that  we  happen  to 
obtain  and  that  seem  worthy  of  recognition. 

It  is  not  to  be  supposed  that  they  will  be  found  constant,  nor  that 
intermediate  forms  will  not  occur.  That  coast  seems  to  be  one  vast 
nursery  for  new  varieties  and  subspecies. 

Some  of  the  odd  forms  are  probably  hybrids  between  distinct 
species,  but  at  present  there  is  no  way  to  determine  this.  Some  of  the 
forms  classed  in  Leptasterias  may  eventually  prove  to  be  the  young 
of  larger  species  of  Asterias  proper,  for  the  latter  all  pass  through 
a  Leptasterias-like  stage  in  the  course  of  their  growth.  It  is  impossi- 
ble, in  many  cases,  to  distinguish  these  young  forms  without  larger 
series  of  intermediate  sizes.  The  only  positive  criterion,  in  many 
cases,  would  be  the  position  of  the  genital  pores  and  the  incubation 
of  the  young,  which  have  not  been  observed  in  many  species. 

LEPTASTERIAS  ( ?)  INEQUALIS  Verrill,  sp.  nov. 
Plate  LXXIII,  figure  2 ;  text-figures  4,  5. 

The  type  is  a  small  five-rayed  starfish  with  depressed,  rapidly 
tapering  and  rather  acute  rays.  Radii,  7  mm.  and  26  mm.;  ratio, 
i :  3.71. 

The  dorsal  ossicles  are  lobate,  numerous,  mostly  flat  and  closely 
united,  leaving  small  papular  areas,  and  not  distinctly  reticulate  in 
arrangement.  The  larger  plates  in  the  median  and  intermediate  rows 
are  convex  and  more  prominent.  The  apical  plates  are  smaller  than 
in  epichlora  of  the  same  size ;  hence  the  rays  are  more  acute.  The 


1 18  VERRILL 

dorsal  spines  are  very  unequal  in  size  and  form.  Those  on  the 
median  radial  rows,  and  in  the  incomplete,  short,  intermediate  series, 
are  much  larger  than  the  rest,  about  as  broad  as  high,  with  round 
capitate  tips.  They  mostly  stand  singly,  but  often  two  to  a  plate. 
Large  numbers  of  very  much  smaller,  unequal,  minute  spinules  are 
thickly  scattered  over  the  surface,  but  not  crowded.  These  stand 
singly  or  in  small  groups  on  the  smaller,  intermediate  ossicles.  Each 
spine  is  surrounded  by  a  circle  of  minor  pedicellariae,  nearly  as  large 
as  the  smaller  spines.  These  small  spines  are  slender,  two  or  three 
times  as  long  as  thick,  terete  or  slightly  clavate,  obtuse,  partially 
concealed  by  the  wreaths  of  pedicellariae. 


X/7 


FIG.  4. 


Leptasterias  inequalis  V.  type,  i,  Portion  of  the  dorsal  surface  showing  spines  of  diverse 
sizes,  wreaths  of  minor  pedicellariae,  and  papular  pores.  X  17.  ii,  The  same;  a,  a',  adam- 
bulacral  spines  and  epispinal  pedicellariae ;  b,  double  row  of  interactinal  spines  with  epispinal 
pedicellariae  and  major  pedicellariae  between  their  bases;  c,  inferomarginal  spines  with 
epispinal  pedicellariae. 

The  disk  is  covered  with  numerous  scattered,  unequal  spines, 
which  do  not  form  very  evident  circles  around  the  madreporic  plate. 
The  latter  is  small,  with  fine  gyri,  and  not  very  far  from  the  center. 

The  upper  marginal  spines  are  decidedly  longer  than  the  median 
and  slightly  tapered,  obtuse  or  subacute ;  they  form  a  regular  single 
row. 

The  inferomarginals  and  interactinals  are  a  little  longer,  more 
slender  and  less  obtuse.  The  inferomarginals  and  peractinals  each 
form  a  single  continuous  row,  the  spines  becoming  shorter  and  more 
obtuse  distally.  The  subactinal  row  extends  to  about  the  middle  of 
the  ray.  The  lateral  channel  between  the  two  marginal  rows  is 
narrow  but  very  distinct. 

The  adambulacral  spines  are  about  as  long  as  the  peractinals,  and 
nearly  as  large.  They  are  terete,  often  a  little  thinner  in  the  middle 
and  slightly  enlarged  toward  the  obtuse  or  subacute  tips;  those 


SHALLOW-WATER   STARFISHES 


119 


towards  the  mouth  are  distinctly  longer,  more  slender,  and  more 
acute.  The  mouth  is  not  much  sunken  and  the  adoral  carinae  are 
short,  composed  of  only  two  contingent  pairs  of  plates,  besides  the 
epiorals.  The  apical  pairs  of  peroral  spines  are  well  developed,  but 
shorter  and  not  stouter  than  the  average  adambulacrals ;  the  side 
spine  is  about  half  as  long,  strongly  divergent,  so  that  those  on 
adjacent  jaws  often  have  their  tips  in  contact  and  thus  might  form 
a  continuous  fence  around  the  oral  area.  The  epiorals  and  adorals 
are  long  and  slender,  tapered,  subacute ;  they  often  bear  both  minor 
and  major  pedicellariae.  The  latter  are  small,  ovate-lanceolate,  sub- 
acute.  Minor  pedicellariae  are  abundant  on  all  the  spines,  forming 
close  wreaths  on  the  dorsals  and  superomarginals,  but  secund  clusters 

* 


FIG.  5. 

Leptasterias  inequalis.  iii,  One  of  the  jaws,  a,  a;  a',  a',  apical  or  peroral  spines; 
a",  lateral  jaw-spines;  e,  the  first  pair  of  epioral  or  suboral  spines;  e' ',  ad  pair;  p,  pedi- 
cellariae;  X  17.  iv,  One  of  the  adoral  spines;  X  17.  v,  A  major  pedicellaria  more  enlarged. 

on  all  the  spines  of  the  lower  surface.  Major  pedicellariae  are  few 
in  number,  small,  ovate  or  ovate-lanceolate. 

The  type  is  from  Orca,  Alaska  (Prof.  W.  R.  Coe,  Harriman 
Expedition). 

This  is,  quite  probably,  the  young  of  a  larger  species,  perhaps  not 
a  Leptasterias,  but  I  am  unable  to  refer  it  to  any  known  to  me.  It 
somewhat  resembles  some  of  the  five-rayed  varieties  of  epichlora, 
but  the  flat,  closely  imbricated  dorsal  ossicles,  without  many  reticula- 
tions, forbid  its  union  with  that  protean  species,  at  present,  for  no 
intermediate  specimens  have  been  found. 

LEPTASTERIAS  LEPTALEA  Verrill,  sp.  nov. 

Plate  xvin,  figure  3  (type). 

A  small  and  very  delicate  five-rayed  species.  Radii,  2.5  mm.  and 
15  mm. ;  ratio,  i :  6.  Rays  terete,  evenly  tapered.  Dorsal  ossicles 
relatively  strong,  thickened,  especially  those  in  the  median  rows. 


I2O  VERRILL 

Dorsal  spines  not  numerous,  conical,  relatively  stout,  usually  stand- 
ing singly,  one  to  an  ossicle,  and  forming  three  or  five  rather  irregu- 
lar rows.  Ocular  plate  relatively  large,  bearing  a  dense  cluster  of 
small  spines.  Minor  pedicellariae  of  unusually  large  size  and  with 
strongly  curved  blades  are  usually  numerous  in  groups  around  the 
dorsal  spines  and  on  the  papular  areas.  Adambulacral  spines  mostly 
two  to  a  plate,  or  alternately  one  and  two  distally,  very  slender, 
terete,  often  slightly  clavate.  Lower  marginal  spines,  which  are  much 
larger,  acute,  conical,  like  the  dorsals,  but  longer,  form  a  single  row 
next  the  adambulacrals,  or  there  may  be  two  to  a  plate,  proximally, 
in  the  larger  specimens.  (The  synactinal  plates  appear  to  be  few 
and  small,  or  altogether  lacking  in  our  examples,  but  may  occur  in 
larger  ones.  Upper  marginal  spines  form  a  single  row  and  resemble 
the  dorsals  in  form  and  size.  Major  pedicellariae  few,  small,  ovate. 

The  ambulacral  feet  are  in  four  rows,  but  the  adjacent  rows  form 
only  a  zigzag  line,  so  that  they  sometimes  appear  almost  biserial. 

This  species  looks  very  much  like  a  Pedicellaster,  not  only  in  size 
and  spinulation,  but  also  in  the  character  of  the  pedicellariae.  The 
arrangement  of  the  ambulacral  feet  in  four  rather  indistinct  rows 
forbids  its  place  in  that  genus,  however. 

Our  specimens  are  evidently  not  full-grown,  but  it  probably  never 
becomes  large,  perhaps  not  more  than  50  mm.  in  diameter. 

Virgin  Bay,  Alaska  (W.  R.  Coe). 

It  is  more  delicate  than  the  Atlantic  L.  tenera,  of  the  same  size, 
but  has  larger  dorsal  spines,  much  fewer  in  number,  and  the  dorsal 
pedicellariae  are  much  larger  and  different  in  form.  The  much 
smaller  number,  larger  size,  and  isolated  position  of  the  dorsal  spines 
indicate  that  it  cannot  be  the  young  of  C.  cribraria  or  L.  hexactis. 
The  young  of  the  latter,  of  similar  size,  are  known  to  me,  and  are 
very  different. 

LEPTASTERIAS  ARCTICA  (Murdoch). 

*  *n  7*. 

Plate  LVI,  figures  I,  2  (young)  ;  plate  LXXI,  figures  I,  2;  plate  ixxn,  figure  i; 

plate  LXJfocin,  figures  2,  20  (details). 

Asterias  arctica  (Murdoch),  Report  of  the  International  Polar  Expedition 
to  Point  Barrow,  Alaska,  under  Lieut  Ray,  1885,  p.  159. 

Mr.  Murdoch's  original  description  is  as  follows : 

"  Rays  five,  rounded  above,  elongated,  tapering  regularly  to  the 

tips.    Radii  as  1 : 3.5.    Disk  small,  its  radius  about  equal  to  width 

of  ray  at  base.     Interambulacral  spines  round  and  slender,  with 

rounded  tips,  usually  two  to  each  plate.     No  small  spines  between 


SHALLOW- WATER   STARFISHES  121 

these  and  the  ventral  spines.  Ventral  spines  (inferomarginals) 
form  a  double  row  of  alternating  spines,  of  which  the  upper  are  the 
smaller  and  the  lower  are  larger  and  stouter  than  the  interambu- 
lacrals.  Lateral  spines  (superomarginals)  rather  slender,  forming 
a  single  row.  No  well  marked  dorsal  row,  though  the  spines  in  the 
middle  of  the  arm  are  rather  the  larger.  The  dorsal  spines  are  short 
and  stout,  with  rounded,  almost  capitate  tips.  The  spines  of  the 
disk  are  rather  smaller  than  those  of  the  arms  and  are  arranged 
irregularly.  The  major  pedicellariae  could  not  be  well  made  out,  but 
appeared  to  be  lanceolate  and  not  numerous.  The  minor  pedicel- 
lariae form  close  wreaths  around  the  spines." 

I  have  examined  the  types  of  this  species,  which  are  preserved  in 
the  U.  S.  National  Museum,  and  also  a  number  of  other  lots  from 
the  same  region. 

The  larger  type-specimen,  dried  from  alcohol  (a),  has  the  radii, 
7  mm.  and  32  mm.;  ratio,  1 : 4.57.  A  smaller  one  (6)  has  the  radii, 
5  mm.  and  23  mm. ;  ratio,  1 : 4.60.  But  it  becomes  considerably 
larger  than  these.  One  of  the  larger  ones  from  Arctic  Alaska  (No. 
1428,  Mus.  Comp.  Zool.,  pi.  LXXI),  has  the  radii  u  mm.  and  44  mm.  ; 
ratio,  i :  4. 

This  species  varies  somewhat  in  the  size  and  number  of  the  dorsal 
and  marginal  spines  and  also  in  the  form  and  size  of  the  adambu- 
lacral  spines,  and  especially  in  the  number  of  minor  pedicellariae. 
The  general  appearance  is  somewhat  like  that  of  L.  compta,  of  the 
New  England  coast,  but  the  forms  of  the  spines  are  coarser  and  the 
pedicellariae  are  different. 

In  most  cases,  including  the  type-specimens  (a  and  b),  there  is  an 
evident,  but  crooked,  median  radial  row  of  distinctly  larger  spines, 
when  dry.  The  other  spines  are  rather  numerous,  irregularly  scat- 
tered, and  reticulated,  but  often  forming  indistinct  longitudinal 
rows,  as  well  as  occasional  short  transverse  series  proximally  on  the 
sides  of  the  rays.  They  are  all  short  and  blunt,  not  minute.  The 
larger  ones  are  about  as  high  as  broad,  somewhat  capitate,  with  finely 
striated  and  evenly  rounded  tips ;  the  smaller  ones  are  more  or  less 
clavate  or  subcapitate.  All  the  dorsal  and  marginal  spines  are 
usually  surrounded  by  dense  clusters  of  very  small  minor  pedicel- 
lariae, which  in  alcoholic  specimens  are  attached  to  a  sheath  that 
usually  rises  to  mid-height  of  the  spine,  and  in  the  case  of  the  infero- 
marginal  spines,  it  frequently  nearly  reaches  the  tip,  and  thus  the 
dense  clusters  of  pedicellariae  often  conceal  the  tips  of  the  spines  and 
are  nearly  or  quite  in  contact  with  each  other.  But  some  specimens, 


122  VERRILL 

of  small  or  moderate  size,  have  only  very  small  clusters  of  pedicel- 
lariae,  and  these  may  be  seen  only  at  the  base  of  the  spines,  espe- 
cially when  dry,  the  sheath  being  rudimentary  or  contracted ;  or  they 
may  be  lacking  on  many  of  the  spines  in  young  specimens. 

The  superomarginal  spines  are  much  like  the  dorsals  in  size  and 
form,  and  therefore  not  easily  distinguished  in  many  specimens,  but 
they  are  usually  slightly  longer.  In  most  specimens  of  average  size 
there  is  a  short  proximal  row  of  more  slender  spines  interpolated 
between  the  upper  and  lower  marginals. 

The  inferomarginal  spines  are  decidedly  longer  and  larger  than  the 
upper  ones,  often  being  twice  as  stout.  They  are  usually  terete, 
scarcely  tapered,  often  slightly  curved  and  obtuse.  Frequently  they 
stand  two  to  a  plate ;  the  upper  ones  are  smaller ;  but  in  many  cases 
they  stand  singly,  in  a  regular  row.  The  larger  specimens  usually 
have  a  short  proximal  row  of  interactinal  spines,  similar  to  the 
inferomarginals.  This  is  true  of  the  larger  of  Mr.  Murdoch's  type- 
specimens.  The  adambulacral  spines  stand  one  or  two  to  a  plate  in 
irregular  alternation.  They  are  nearly  as  long  as  the  inferomarginals, 
usually  much  more  slender,  terete  and  blunt,  sometimes  slightly 
clavate,  but  in  some  cases  they  are  stouter  than  usual  and  nearly  as 
large  as  the  inferomarginals.  They  usually  bear  clusters  of  minor 
pedicellariae,  and  many  isolated  major  pedicellariae. 

The  apical  peroral  spines  are  usually  rather  large  and  strong, 
being  considerably  stouter  than  the  adorals.  The  epiorals  are  also 
rather  stout  and  distinctly  longer  than  the  adorals. 

Major  pedicellariae  are  sometimes  few  in  number,  but  usually  there 
are  many  of  rather  small  size  attached  within  the  margin  of  the 
grooves,  and  some  of  larger  sizes  on  and  between  the  adambulacral 
spines  and  oral  spines.  Some  of  the  larger  kind  also  occur  on  the 
interradial  areas  and  lateral  channels.  They  are  mostly  compressed, 
ovate-lanceolate  or  oblong-lanceolate,  with  acuminate  tips.  The 
larger  ones  are  often  as  stout  as  the  adjacent  adambulacral  spines, 
and  these  usually  have  dentate  tips. 

Mr.  Murdoch  described  the  same  specimens,  as  preserved  in 
alcohol,  hence  he  overlooked  some  of  the  details,  obvious  when  dry. 

Two  young  specimens,  which  I  refer  doubtfully  to  this  species, 
were  collected  by  Dr.  W.  H.  Dall  on  the  coast  of  East  Siberia,  at 
East  Cape  (U.  S.  Nat.  Mus.,  No.  6031). 

Others,  similar,  were  collected  in  Bering  Strait  by  Robt.  White 
(No.  16591.  PI.  LVI.  figs,  i,  2.)  The  radii  of  the  largest  are 
3.5  mm.  and  n  mm.  The  dorsal  spines  are  small,  clavate,  arranged 


SHALLOW-WATER   STARFISHES  123 

in  three  radial  rows,  those  in  the  median  one  largest.  The  two 
marginal  rows  are  regular  and  distinct,  the  spines  being  longer, 
regularly  spaced,  one  to  each  plate.  Actinals  mostly  lacking.  Adam- 
bulacral  spines  slender,  subdiplacanthid.  Minor  pedicellariae  are 
present  in  small  clusters  on  most  of  the  spines,  but  form  complete 
wreaths  on  the  upper  marginals.  One  of  these  young  specimens  was 
labelled,  by  Dr.  Dall,  as  white  when  living. 

Arctic  Ocean,  off  Point  Franklin,  in  thirteen  and  one-half 
fathoms,  sand  (coll.,  Murdoch,  Point  Barrow  Expedition),  types 
(three),  No.  7625,  U.  S.  National  Museum;  Arctic  Ocean,  ten 
fathoms  (Dr.  W.  H.  Dall),  No.  3622  (1669),  U.  S.  National 
Museum;  Bering  Straits  (Robt.  White),  No.  16591;  East  Siberia, 
Nos.  6031  and  16584  (young)  (W.  H.  Dall). 

Several  young  specimens  (Nos.  16584)  from  ten  to  fifteen  fath- 
oms, Plover  Bay,  Siberia,  probably  of  this  species,  are  nearly  or  quite  '  ^^x 
monacanthid.  The  larger  have  radii  5  mm.  and  15  mm.  The  dorsal 
spines  are  unequal,  not  crowded,  the  larger  are  short,  capitate;  all 
are  surrounded  by  pedicellariae  in  wreaths,  unusually  abundant  for  so 
young  specimens.  Papulae  are  mostly  isolated ;  dermis  thick. 

LEPTASTERIAS  COEI  Verrill,  sp.  nov. 
Plate  ix,  figure  i;  plate  xvii,  figures  I,  2  (types). 

A  slender  species,  usually  six-rayed.  Disk  small;  rays  long  and 
neatly  rounded,  tapering  gradually.  Radii,  5  mm.  and  30  mm.; 
ratio  i :  6,  sometimes  less. 

The  dorsal  spinules  form  three  pretty  regular  primary  rows,  with 
many  smaller  ones  scattered  between  irregularly,  but  not  crowdedly, 
and  not  clustered.  Usually  the  spines  stand  singly,  one  to  a  plate; 
the  median  radial  ones  are  larger  and  form  a  pretty  regular  row. 
All  are  cylindrical  or  slightly  clavate,  of  moderate  length,  with 
rounded  tips,  and  are  surrounded  by  small  dense  wreaths  of  minor 
pedicellariae,  which  are  nearly  in  contact  in  alcoholic  specimens. 

The  upper  marginals  are  a  trifle  longer  and  form  a  regular  row, 
one  to  a  plate.  The  lower  marginals  are  rather  longer,  terete, 
mostly  one  to  a  plate,  but  sometimes  with  a  second  smaller  one 
proximally.  An  imperfect  row  of  somewhat  smaller  synactinal 
spines  proximally.  The  adambulacral  spines  are  mostly  in  two 
equal  rows,  but  some  of  the  plates  bear  only  one  spine ;  these  spines 
are  slender,  terete,  tapered,  subacute,  nearly  as  long  as  the  lower 
marginals,  but  much  more  slender;  they  bear  small  groups  of  minor 


124  VERRILL 

pedicellariae.  A  few  small,  long-ovate,  compressed  major  pedicel- 
lariae  occur  on  the  actinal  interradial  areas  and  along  the  edges  of  the 
grooves.  The  papulae  stand  singly  or  in  groups  of  two  to  four. 
Madreporic  plate  small,  prominent,  with  few  gyri,  pale  orange  in 
alcohol. 

A  smaller  specimen  (pi.  ix,  fig.  I ;  pi.  xvn,  fig.  2),  taken  at  the 
same  place,  was  also  preserved  in  alcohol.  The  dorsal  spines  already 
form  five  distinct  rows,  with  others  between  them ;  the  median  row  is 
conspicuous  and  its  spines  much  larger  than  the  others,  with  rounded 
tips.  Wreaths  of  minor  pedicellariae  are  much  smaller  and  not  in 
contact  generally.  Lateral  and  ventral  spines  longer  and  in  very 
regular  rows.  Madreporic  plate  is  pale  orange  in  alcohol.  In  life 
the  color  was  uniform  iron-rust  color  (Coe). 

The  types  described  are  from  Berg  Bay,  Alaska,  in  ten  fathoms 
(W.  R.  Coe,  Harriman  Expedition,  June  10,  1899). 

This  species  bears  a  general  resemblance  to  L.  tenera  (St.)  of  the 
New  England  coast,  but  the  dorsal  and  marginal  spines  in  the  latter 
are  far  more  numerous,  much  smaller,  and  acute ;  it  very  rarely  has 
six  arms,  and  its  dorsal  ossicles  are  much  more  numerous.  Typical 
specimens  of  L.  mulleri  from  Greenland  are  still  more  closely  allied, 
but  the  latter  is  a  five-rayed  species,  and  also  has  more  slender  spines 
(though  not  so  slender  as  in  L.  tenera},  and  has  fewer  pedicellariae, 
somewhat  different  in  form. 

LEPTASTERIAS  MACOUNI  Verrill,  sp.  nov. 

Rays  six,  elongated,  slender,  rounded  and  evenly  tapered.  Radii, 
8  mm.  and  41  mm. ;  ratios,  1 :  5.  Dorsal  ossicles  thick  but  small,  with 
large  papular  pores  between  them;  median  row  thicker,  forming  a 
slight  carina;  others  reticulate.  Dorsal  spines  slender,  small,  scat- 
tered or  in  several  obscure  rows,  except  the  median  radial  row,  which 
is  distinct,  consisting  of  small  clavate  spines,  mostly  standing  two  to 
a  plate,  in  nearly  simple  lines.  Similar  spines  cover  the  disk.  The 
larger  intermediate  dorsal  spines  are  rather  smaller,  slender,  tapered, 
subacute,  mostly  one  to  a  plate,  arranged  somewhat  in  quincunx,  or 
in  about  three  irregular  rows  on  each  side,  the  larger  papular  areas 
forming  oblique  rows  of  three  on  each  side.  Some  minute  spines 
stand  on  the  transverse  ossicles. 

Superomarginal  spines  distinct,  larger  and  longer  than  dorsals, 
obtuse  or  a  little  clavate,  forming  a  simple  regular  row.  Infero- 
marginal  spines  are  of  similar  length  or  a  little  longer,  subacute, 
mostly  two  to  a  plate,  forming  two  alternating  rows. 


SHALLOW- WATER   STARFISHES  12$ 

Interactinal  spines  are  similar  to  the  inferomarginals  and  form 
three  simple  oblique  rows  proximally,  one  spine  to  a  plate,  there 
being  three  rows  of  small,  thick  interactinal  ossicles  separated  by 
rows  of  small  papular  pores.  The  peractinal  row  reaches  nearly  to 
the  tip  of  the  ray ;  the  next  one  reaches  to  about  the  middle ;  while  4  -K-*-^  . 
the  third  is  confined  to  the  basal  part  of  the  ray.  All  these  rows  bend 
upward  pretty  regularly  proximally. 

Adambulacral  spines  slender,  tapered,  obtuse,  much  smaller  and 
shorter  than  the  interactinals,  subdiplacanthid,  part  of  the  plates 
bearing  one,  and  part  two  spines  in  irregular  alternations.  Oral 
spines  have  been  destroyed  in  the  type. 

Small  wreaths  of  minute  minor  pedicellariae  surround  most  of  the 
dorsal  spines ;  major  pedicellariae  few  and  small.  Madreporite  small, 
fine-grained,  surrounded  by  a  circle  of  small  clavate  spinules. 

The  type  was  taken  at  Departure  Bay,  Vancouver  Island,  on  rocks 
at  low  tide  by  Prof.  John  Macoun  and  party,  of  the  Canadian  Geo- 
logical Survey,  1909  (No.  43).  It  is  dedicated  to  Prof.  John  Ma- 
coun. I  have  seen  no  others. 

This  species  most  resembles  L.  hexactis  in  general  appearance,  but 

"*"*        -^r-if 

the  latter  has  the  dorsal  spines  clustered  and  has  fewer  rows  of 
actinal  plates  and  spines.  It  may  eventually  prove  to  be  a  variety  of 
hexactis,  when  a  larger  series  can  be  studied.  I  have  seen  no  inter- 
mediate specimens. 

LEPTASTERIAS  VANCOUVERI  (Perrier). 

Asterias  vancouveri  PERKIER,  Arch,  de  Zool.  Exper.,  iv,  p.  328,  1875.    Bell, 
Trans.  Zool.  Soc.  London,  pp.  496,  504,  1881.     ?  —  A  hexactis  Stimpson. 

No  specimen  perfectly  agreeing  with  this  small  diplacanthid 
species  has  come  under  my  observation. 

According  to  the  imperfect  description  of  M.  Perrier,  it  somewhat 
resembles,  in  form  and  ornamentation,  Asterias  (Stichaster)  poly- 
plax.  Arms  six  or  seven.  Adambulacral  spines  long,  pointed,  two 
upon  each  plate,  but  not  in  two  regular  rows,  the  rows  irregularly  al- 
ternating; each  bears  on  the  middle  one  or  two  small  major  pedicel- 
larise.  Outside  of  these  are  two  rows  of  pointed  ventral  spines,  larger 
than  the  adambulacrals  and  surrounded  by  a  few  minor  pedicel- 
larise.  Ossicles  of  the  dorsal  skeleton  much  as  in  ordinary  species  of 
Asterias.  On  the  back  are  five  ranges  of  groups  of  small  obtuse 
spines,  each  ossicle  bearing  a  group  of  three  or  four  spines ;  other 
spines  are  borne  on  the  transverse  connecting  ossicles,  so  that  the 
groups  of  spines  appear  confluent  and  form  a  vague  reticulation. 


VERRILL 

Madreporic  plate  single.  Diameter  of  disk,  10  mm.;  distance 
between  ends  of  the  two  arms,  75  mm. 

Vancouver  Island  (British  Museum). 

This  small  species,  to  judge  from  the  brief  description,  should  be 
closely  allied  to,  or  identical  with,  L.  hexactis,  of  earlier  date.  The 
description,  at  least,  would  apply  to  that  species.  The  types  of  the 
latter,  moreover,  were  from  the  same  region  (Puget  Sound).  No 
other  similar  six-rayed  species  has  been  found  by  me  in  any  of  the 
large  collections  recently  received  from  Vancouver  Island. 

It  must  also  resemble  some  of  the  more  slender  six-rayed  varieties 
of  L.  epichlora,  but  the  latter  does  not  often  occur  in  my  Vancouver 
Island  collections,  and  has  much  more  numerous  actinal  spines. 

LEPTASTERIAS  HEXACTIS  (Stimpson). 
Plate  xxv,  figures  7,  8  (type). 

Asterias  hexactis  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vm,  p.  272,  1862. 
Verrill,  Trans.  Conn.  Acad.  Sciences,  i,  p.  326,  1867.    Bell,  1881,  p.  495. 

This  is  a  small  six-rayed  starfish,  much  like  L.  aqualis,  but  with 
the  dorsal  spinules  longer,  slender,  less  numerous,  and  not  so  closely 
clustered.  The  rays  are  usually  longer  and  more  tapered. 

Dr.  Stimpson's  description  was  as  follows : 

"  Rays  six,  depressed  or  rounded,  and  more  or  less  tapering.  Disc 
large.  Proportion  of  the  diameters,  i :  4.  Ambulacral  spines,  sub- 
equal,  in  two  regular  rows  towards  the  disc,  two  to  each  plate; — 
form  cylindrical,  obtuse,  sometimes  a  little  clavate,  with  a  few  pedi- 
cellariae  of  both  kinds  on  their  outer  side  at  the  middle.  On  the  lat- 
ero-inferior  side  of  the  ray  there  are  four  longitudinal  rows  of  spines, 
separated  from  the  dorsal  spines  by  a  more  or  less  well  marked 
channel.  These  spines  are  scarce  thicker  than  the  ambulacrals,  but 
are  longer,  and  have  small  clusters  of  minor  pedicellariae  at  their 
outer  bases.  In  some  specimens  the  lateral  spines  are  distinct  from 
the  ventrals,  being  separated  from  them  by  a  channel,  and  forming 
a  crowded  row  of  confluent  clusters  like  the  dorsals.  Dorsal  spines 
small  and  numerous,  in  little  heaps,  which,  being  confluent  in  a  longi- 
tudinal direction,  form  three  or  five  (according  to  the  distance  from 
the  disc)  rows,  separated  from  each  other  by  corresponding  rows  of 
papuliferous  depressions.  These  spines  in  some  specimens,  however, 
are  fewer,  and  do  not  form  heaps.  On  the  disc  they  are  arranged 
after  a  reticulating  pattern.  The  spines  are  capitate,  and  sparsely 
surrounded  by  minor  pedicellariae.  All  rise  to  about  the  same  height, 
thus  giving  an  evenness  to  the  outline  as  seen  in  a  side  view.  The 


SHALLOW-WATER   STARFISHES  127 

major  pedicellariae  are  few,  and  formed  on  the  labial  spines,  or  rarely 
a  single  one  on  the  side  of  the  ray ; — they  are  more  or  less  pointed, 
about  one-fortieth  of  an  inch  long,  and  twice  as  long  as  broad. 
Dorsal  papulae  in  small  groups.  Ventral  papulae  mostly  single,  and 
curving  upward  or  outward.  Diameter,  one  and  three-fourths  <— -,  , 
inch.  A  variety  occurs  with  more  slender  and  tapering  rays. 

"  It  is  smaller  than  A.  Camtschatica  Brandt,  and  has  longer  arms,   «T>  I 
etc.  / ' 

"  Habitat,  Puget  Sound.    North  West  Boundary  Commission.    Dr.    l^*i*t? > ($t IK 
C  B.  Kennedy."  ^.  C&f. 

A  good  dry  specimen  in  the  Yale  Museum,  from  Monterey 
(Stevens),  seems  to  agree  pretty  closely  with  the  type.  The  radii  -^ 
are  5  mm.  and  22  mm.;  ratio,  about  1:4.5.  Rays  six,  slender, 
terete,  tapered  to  acute  tips.  The  dorsal  spines,  which  mostly  stand 
singly,  sometimes  two  or  three  together,  do  not  form  definite  rows. 
They  are  numerous,  but  not  crowded,  slender,  slightly  clavate,  or 
subacute;  they  are  surrounded  by  small  wreaths  of  minor  pedicel- 
lariae,  of  relatively  large  size,  which,  in  alcoholic  specimens,  fill  up 
the  spaces  between  the  spines  and  give  the  surface  the  same  even 
appearance  as  if  the  spines  were  more  crowded.  The  upper  mar- 
ginal spines,  which  are  nearly  like  the  dorsals  in  size,  though  rather 
longer,  form  a  pretty  regular  row;  two  often  stand  on  one  plate. 
The  lower  marginals  are  at  least  twice  as  large  and  long,  and  form 
a  very  distinct  row,  but  two  often  stand  on  a  plate ;  they  are  terete, 
tapered,  subacute.  The  few  synactinal  spines,  when  present,  are 
similar.  Adambulacrals  are  nearly  as  long  as  the  marginals,  but  are 
more  slender  and  scarcely  taper;  they  mostly  stand  two  to  a  plate, 
divergently,  and  form  two  regular  rows ;  they  become  distinctly 
longer  near  the  mouth.  The  tip  of  each  jaw  bears  two  terminal 
longer  and  stouter  spines,  larger  than  the  marginals,  and  a  much 
smaller  one  external  to  these,  on  each  side. 

Major  pedicellariae  of  rather  small  size  are  borne,  mostly  singly, 
on  many  of  the  adambulacral  spines,  and  much  smaller  ones  on  the 
inner  margins  of  the  grooves.    They  are  compressed,  long-ovate  or     . 
lanceolate,  subacute.     They  are  larger,  longer,  and  less  triangular 
than  those  of  L.  ceqwAls. 

A  young  specimen,  25  mm.  in  diameter,  from  Kadiak,  Alaska 
(Harriman  Expedition),  appears  to  belong  to  this  species.  The  six 
rays  are  very  slender,  and  rather  closely  covered  with  very  small 
obtuse  and  capitate  spines,  more  or  less  areolated  on  the  central  parts. 
Adambulacral  spines  very  slender,  mostly  in  two  rows.  Its  aspect  is 
like  that  of  a  Pedicellaster. 


128  VERRILL 

Dr.  R.  Rathbun  has  furnished,  for  reproduction  here  (pi.  xxv, 
figs.  7,  8),  photographs  of  the  type,  now  in  the  U.  S.  National 
Museum. 

San  Francisco,  California,  to  Sitka.  A  common  littoral  and  shal- 
low-water species.  At  Monterey  it  occurs  associated  with  L.  aquatis. 

I  have  also  studied  numerous  specimens  from  Puget  Sound  and 
Queen  Charlotte  Islands,  sent  by  the  Geological  Survey  of  Canada. 

This  species  has  often  been  confounded  with  L.  aqualis  in  col- 
lections. Though  closely  related,  it  can  be  distinguished  by  its  dif- 
ferent spinulation  and  distinct  form  of  pedicellariae. 

LEPTASTERIAS  ^EQUALIS  (Stimpson). 

Plate  xvi,  figure  8;  plate  xvm,  figures  i,  2;  plate  xxv,  figures  5,  6  (type) ; 
plate  LVI,  figure  5  (var.). 

Asterias  eequalis  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vni,  p.  273,  1862. 
Verrill,  op.  tit,  p.  327,  1867. 

As  usually  found,  this  is  a  small,  six-rayed  species.  Disk  small ; 
rays  rather  short.  Dorsal  spines  small,  clavate  or  capitate,  sulcate, 
and  rough  or  spinulose  at  tip,  very  numerous,  covering  the  whole 
upper  surface  nearly  evenly,  but  sometimes  forming  indistinct  median 
radial  rows.  A  group  or  row  of  spines  stands  on  each  of  the  ossicles, 
which  are  small  and  numerous;  they  are  rather  closely  imbricated, 
but  leave  papular  areas,  often  as  wide  as  the  ossicles.  Median 
ossicles  larger,  with  rounded  lobes,  the  proximal  lobe  longer  and 
broader,  imbricated.  Marginal  and  actinal  spines  more  elongated, 
forming  several  longitudinal  rows,  slender,  subacute.  Papular  areas, 
on  the  upper  side,  form  longitudinal  rows ;  the  papulae  stand  singly  or 
sometimes  two  or  three  together. 

A  good  characteristic  specimen  of  the  California  form  (var. 
cequalis)  from  Monterey,  the  type  locality,  has  the  radii  6.5  mm.  and 
20  mm. ;  ratio,  about  1 : 3.5.  The  rays  are  round  and  plump,  taper- 
ing gradually. 

The  dorsal  spines  are  very  numerous,  small,  nearly  equal  and 
even,  mostly  capitate  or  clavate,  with  sulcate  tips;  they  stand  in 
groups  of  three  to  eight  or  more.  Some  of  the  groups  are  circular 
with  a  central  spine,  but  others  surround  the  papular  pores,  which 
have  an  evident  longitudinal  arrangement.  The  upper  marginal 
plates  form  a  crowded  row,  two  to  four  spines  standing  together  on 
most  of  the  plates,  while  one  or  two  usually  stand  on  the  descend- 
ing lobe  or  connecting  ossicle,  proximally ;  these  spines  are  slightly 
longer  than  the  dorsals,  and  mostly  bent  upward,  but  are  otherwise 


SHALLOW- WATER   STARFISHES  129 

similar.  The  lower  marginal  spines  form  a  double  row,  two 
divergent  spines  usually  on  each  plate;  these  spines  are  distinctly 
larger  and  longer  than  the  dorsals,  mostly  compressed  and  somewhat 
clavate  at  the  tip,  and  usually  bent  outward  and  upward.  A  single 
row  of  synactinal  spines  of  similar  size  and  shape  usually  occurs  on 
the  proximal  part  of  the  rays.  Adambulacral  spines  are  nearly  as 
long  as  the  actinals  and  about  half  as  thick,  terete,  obtuse.  On  the 
basal  half  of  the  rays  they  stand  two  on  a  plate,  forming  two 
divergent  rows ;  distally  they  are  irregularly  alternated,  one  and  two ; 
those  near  the  mouth  are  longer.  The  tip  of  the  jaw  bears  four 
divergent  spines,  the  two  central  stouter  but  not  longer  than  the 
adambulacrals,  the  other  two  much  smaller  and  about  half  as  long. 

Major  pedicellariae  of  very  small  size  occur  singly  on  many  of  the 
adambulacral  spines;  these  are  short-ovate  or  triangular-ovate  with 
acute  tips.  They  are  smaller,  shorter,  and  more  triangular  than 
those  of  L.  hexactis.  Smaller  ones  also  occur  on  the  inner  edges  of 
the  grooves.  Major  pedicellariae  of  very  small  size  occur  singly  or 
in  small  clusters  on  most  of  the  dorsal  and  lateral  spines,  and  in 
somewhat  larger  groups  on  the  ventral  ones.  They  also  occur  singly,  ^ 
between  the  spines,  on  the  back.  A  o  •  »  3  /T« 

Dr.  R.  Rathbun  has  furnished  the  photographs  of  the  original  C+4*  A.^.Tfl 
type  of  this  species,  which  is  still  in  the  U.  S.  National  Museum,  j-g  |n  *+***•  *  I 
(PI.  xxv,  figs.  5.  6.)  -^  ••  tfcii 

It  agrees  very  well  with  several  specimens  in  the  Museum  of  Yale 
University,  from  the  same  locality.  Stimpson's  description  is  as 
follows : 

"  Rays  six,  rather  slender  and  much  tapering.  Proportion  of  the 
diameters,  i :  3.5.  This  species  has  a  general  resemblance  to 
A.  hexactis  in  shape,  etc.,  but  differs  in  the  character  of  its  spines, 
particularly  the  very  numerous  dorsals,  which  are  uniform  in  size, 
and  shorter  and  more  crowded,  giving  to  the  back  in  a  much  greater 
degree  that  general  evenness  of  surface  which  is  characteristic  of  the 
Stichasters  and  Cribrellce  \Hewr\c\a ] .  These  spines  are  deeply  stri- 
ated or  radiated  on  their  flattened  heads,  each  showing  eight  or  nine 
ridges.  On  the  side  of  the  ray  there  are  two  or  three  rows  of  longer 
spines,  also  striated.  The  ambulacral  spines  are  for  the  most  part  ar- 
ranged alternately  one  and  two  to  each  plate,  but  there  are  two  to 
each  plate  near  the  disc.  There  are  minor  pedicellariae  about  all  the 
spines,  as  in  the  preceding  species  [A.  hexactis],  but  they  are  much 
less  numerous.  We  can  discover  no  major  pedicellariae  excepting  an 
occasional  small  pointed  one  in  the  ambulacral  furrows.  They  would 
xo 


I3O  VERRILL 

perhaps  be  found  on  the  sides  of  the  ray  in  specimens  more  perfect 
than  those  we  possess.  The  papulae  stand  singly  or  in  groups  of 
three  or  four,  arranged  in  indistinct  longitudinal  rows.  Diameter, 
one  inch  and  a  half. 

"  Habitat,  Monterey,  California.    A.  S.  Taylor." 

Common  on  the  coast  of  California.  Ranges  from  San  Diego 
north  to  Puget  Sound  and  Vancouver  Island.  Monterey  (Stimpson; 
Stevens;  R.  E.  C.  Stearns). 

This  species  is  not  a  typical  Leptasterias.  It  differs  from  most 
species  of  that  genus  in  the  apparent  longitudinal  or  "  stichasterial  " 
arrangement  of  dorso-lateral  plates,  and  in  having  the  minute  spines 
closely  clustered  on  the  plates,  much  as  in  Stephanasterias  and  in 
Henricia,  so  that  its  appearance  is  superficially  like  one  of  the  Stichas- 
terinse.  However,  its  dorso-lateral  plates  are  not  so  wide,  nor  so 
closely  imbricated,  as  in  that  group,  nor  even  so  much  so  as  in  Sten- 
asterias.  Its  genital  pores  and  reproduction  are  not  known. 

This  species  varies  considerably  in  its  form  and  spinulation.  I 
have  examined  numerous  specimens  from  Monterey,  the  type 
locality,  collected  by  several  different  persons.  It  seems  to  be  very 
abundant  on  that  part  of  the  coast,  between  tides. 

Owing  to  its  variability  it  may  be  divided  into  several  varieties 
or  local  races,  though  these  doubtless  intergrade  at  intermediate 
stations. 

The  rather  dwarf  littoral  form  from  California  is  certainly  the 
original  or  type-form,  and  should  therefore  be  designated  as  variety 
csqualis,  if  varieties  be  recognized. 

LEPTASTERIAS  JEQUALIS  Var.  COMPACTA  Verrill,  nov. 

Plate  LVI,  figure  5. 

The  type  of  this  form  is  much  larger  and  more  fully  developed 
than  the  ordinary  littoral  varieties.  The  radii  are  15  mm.  and 
45  mm. ;  ratio,  about  i :  3. 

The  six  arms  are  evenly  rounded  above  and  densely  covered  with 
small  capitate  spines  standing  in  large  clusters  on  each  ossicle.  They 
form  a  distinct,  wide,  median  band,  and  two  other  similar  bands  on 
each  side  are  defined  by  the  rows  of  papular  pores.  The  dorsal 
ossicles  are  in  five  rows,  strong,  elevated  in  the  middle,  closely  united 
in  rows  and  somewhat  imbricated,  especially  in  the  median  row.  A 
close  circle  of  small  capitate  spines,  like  the  dorsals,  surrounds  the 
madreporic  plate,  which  has  numerous  fine  irregular  gyri.  The 


SHALLOW- WATER   STARFISHES  131 

upper  marginal  spines  are  numerous,  similar  to  the  dorsals,  but 
rather  longer ;  they  stand  in  oblique  clusters  on  each  ossicle.  Besides 
the  main  row,  there  are  two  to  four  similar  spines  on  the  descending 
apophysis,  and  on  a  small  connective  ossicle,  sometimes  present, 
forming  an  interpolated  series  between  the  upper  and  lower  mar- 
ginals on  the  basal  half  of  the  rays. 

The  longer  ventral  spines  form  three  main  rows,  besides  a  few 
irregular  smaller  ones. 

The  inferomarginal  plates  are  closely  united  and  each  usually 
bears  two  cylindrical  or  slightly  clavate,  blunt  spines,  decidedly 
longer  and  larger  than  the  dorsals ;  or  sometimes  three  near  the  base 
of  the  rays,  besides  one  or  two  smaller  ones.  The  actinal  spines 
form  a  single  row,  one  to  a  plate,  and  do  not  extend  to  the  tips  of  the 
rays ;  they  are  sometimes  lacking. 

The  adambulacral  spines  stand  two  on  a  plate,  but  not  in  regular 
lines ;  they  are  shorter  and  more  slender  than  the  actinals,  but  similar 
in  form. 

The  superomarginal  ossicles  are  stout,  thick,  and  imbricated, 
strongly  four-lobed.  The  upper  lobe  is  large  and  runs  slightly 
obliquely  outward ;  the  large  lower  lobe  or  apophysis  is  elevated  in 
the  middle,  where  it  bears  several  intermediate  spines,  or  it  may  be 
overlapped  partially  by  a  small  interpolated  ossicle.  There  is  a  row 
of  small  papular  areas  between  the  apophyses.  The  inferomarginals 
are  also  stout,  but  smaller,  with  four  short  lobes.  The  actinal  plates, 
near  the  base  of  the  rays,  are  smaller,  irregularly  elliptical,  and 
closely  united  to  the  adambulacrals.  The  apex  of  the  jaws  bears  a 
pair  of  stout  peroral  spines  and  a  pair  of  divergent  lateral  ones,  about 
half  as  long.  On  the  outer  surface  of  the  jaw  there  are  usually  one 
or  two  pairs  of  epiorals  that  are  longer  and  more  tapered  than  the 
following  ones.  The  ambulacral  grooves  are  unusually  wide ;  the 
plates  not  very  crowded;  the  pores  large,  not  crowded.  Dorsal 
papulae  few,  in  small  groups  or  isolated. 

Minor  pedicellariae  are  very  small  and  occur  on  nearly  all  the 
spines  in  small  groups  or  singly.  Major  pedicellariae,  of  very  small 
size,  acute-triangular  in  form,  occur  along  the  inner  edges  of  the 
ambulacral  grooves. 

Pacific  Grove,  Monterey,  California  (Prof.  W.  R.  Coe)  ;  British 
Columbia. 

This  may  be  regarded  as  the  most  perfectly  developed  variety  of 
L.  aqualis.  Its  characters  are  probably  in  part  due  to  its  large 
size,  but  I  have  seen  others  of  the  same  variety  not  over  an  inch 
in  diameter. 


132  VERRILL 

LEPTASTERIAS  yEQUALIS  Var.  NANA  Verrill,  nov. 

This  name  is  proposed  for  a  common,  rather  dwarfed  variety, 
having  short  obtuse  rays,  rather  densely  covered  with  small,  short, 
nearly  even,  obtuse  or  clavate  spinules. 

The  inferomarginal  and  actinal  spines  are  crowded  together  and 
form  about  four  close  rows. 

Common  on  the  California  coasts;  Gulf  of  Georgia. 

LEPTASTERIAS  ^EQUALIS  Var.  CONCINNA  Verrill,  nov. 

This  name  is  proposed  for  a  larger  form  of  this  species,  in  which 
the  dorsal  spinules  form  large  groups  on  the  ossicles,  and  these  lie 
in  pretty  regular  longitudinal  rows,  three  or  five,  separated  by  very 
distinct  rows  of  papular  areas.  The  larger  marginal  spines  also  form 
very  regular  rows.  tftc^  '^/"^ 

The  type  is  from  Monterey  Bay,  California. 

LEPTASTERIAS  EPICHLORA  (  Brandt)  . 

Plate  xvr,  figures  1-6  (varieties)  ;  plate  xxvin,  figures  i,  2  (variety)  ;  plate 
LXXXV,  figures  i-id  (details),  figures  2-2e  (young). 

Asterias  epichlora  BRANDT,  Prod.  Descr.  Anim.  Mertens,  p.  270,  1835.     (Five- 

rayed  form.    Description  poor.)     ?  Stimpson,  Journ.  Boston  Soc.  Nat. 
i  Hist.,  vi,  p.  528   (excel,  synonyms).     Perrier,  Revis.  Stell.,  p.  331    (no 

description). 
?  Asterias  camtschatica  BRANDT,  Prodromus,  p.  270,  1835  J  Middendorff  Reise, 

ii,  p.  32,  1851.     (Six-rayed  form.) 
Asterias  saanichensis  DE   LORIOL,   Mem.   Soc.   Phys.   et   Hist.   Nat.   Geneve, 

x*xn,  p.  23,  pi.  ii,  figs.  3-3e/,  4,  5,  1897.     (Five-rayed  variety.) 

A  rather  small  starfish,  commonly  60  mm.  to  80  mm.  in  diameter, 
sometimes  150  mm.;  usually  greenish  or  olive  on  the  dorsal  surface 
*n  ^e>  generally  six-rayed,  but  not  infrequently  five-rayed.  The 


rays  are  rather  short  and  plump,  covered  above  with  numerous 
small,  short,  crowded  spines,  which  are  usually  unequal  in  size  and 
mostly  capitate,  but  sometimes  most  of  them  are  more  slender  and 
clavate.  The  dorsal  spines  generally  form  an  irregularly  reticulate 
or  areolate  pattern,  but  are  sometimes  acervate,  forming  nodular 
groups  in  one  variety.  Sometimes  they  are  more  unequal  in  size, 
the  larger  ones  standing  in  one  or  more  radial  rows,  and  not 
acervate.  The  median  rows  may  be  distinct  or  indistinct.  More 
rarely  the  spines  are  all  nearly  equal  and  of  the  smaller  sort  ;  some- 
times they  are  subequal,  crowded,  and  all  of  the  larger  sort. 


SHALLOW- WATER   STARFISHES  133 

_) 

Superomarginals  similar  to  the  dorsals.  Inferomarginal  spines  a 
little  longer,  usually  in  two  or  three  rows.  Peractinals  in  one  or 
two  rows.  Adambulacral  spines  diplacanthid. 

Minor  pedicellariae  usually  numerous.  Frequently,  but  not  in  all 
varieties,  nor  in  the  young,  there  are  a  few  very  large,  stout,  stone- 
hammer-shaped  or  wedge-shaped  serrate  dermal  pedicellariae  on  the 
sides  of  the  rays  or  on  the  interradial  spaces  beneath. 

Its  known  distribution  is  from  Vancouver  Island  to  Yakutat  and 
Dutch   Harbor,   and   St.   Paul's   Island,  Alaska.     It   is  abundant    \  ft-^  *  -wW-t* 
between  tides  on  rocky  shores  and  in  shallow  water. 

Owing  to  its  great  variability,  especially  in  the  dorsal  spines,  it 
seems  to  me  desirable  to  distinguish  several  named  varieties,  which 
are  described  below.  The  varieties  often  appear  as  different  as  dis- 
tinct species. 

The  oral  spines  in  this  species  are  more  feebly  developed  than  in 
most  of  the  allied  forms,  but  vary  considerably.  The  peroral  spines 
usually  consist  of  only  two  rather  short  apical  ones  on  each  jaw, 
usually  distinctly  stouter  than  the  adorals,  usually  straight,  but 
sometimes  curved  a  little  toward  each  other,  so  that  their  tips  are 
convergent.  The  small  side-spines  are  generally  lacking  and  replaced 
by  a  pedicellaria,  but  in  some  of  the  larger  specimens  it  is  present 
on  some  of  the  jaws  and  not  on  others;  when  present  it  is  small  and 
short,  acute,  not  more  than  one-third  or  one-fourth  the  length  of  the 
apical  pair,  but  it  is  variable. 

The  epioral  spines  are  rather  longer  and  more  slender  than  the 
perorals,  and  like  the  adorals  in  form,  though  a  trifle  longer.  It  often 
happens  that  one  of  a  pair  is  lacking  on  some  of  the  jaws  and  then 
the  single  spine  stands  nearly  in  the  middle  of  the  jaw.  The  adoral 
carina  consists  of  three  contingent  pairs  of  adoral  plates,  besides 
the  epioral ;  their  single  spines  are  decidedly  more  slender  than  those 
more  distal,  and  are  a  little  longer.  Double  rows  of  spines  usually 
commence  on  the  fifth  to  the  eighth  plate. 

The  normal  young  of  this  species,  when  10  mm.  to  15  mm.  in 
diameter,  have  a  rather  openly  reticulated  dorsal  skeleton,  composed 
mostly  of  deeply  lobed  ossicles,  with  three  or  four  unequal  lobes. 
When  20  mm.  to  25  mm.  in  diameter,  they  usually  have  the  char- 
acteristic spinulation  of  the  adults,  though  the  spines  are  smaller  and 
much  less  numerous,  and  the  median  radial  row  is  very  distinct  and 
pretty  regular.  The  papulae  are  few  and  mostly  stand  singly.  The 
madreporic  plate  is  small,  with  but  few  gyri.  The  superomarginal 
and  inferomarginal  plates  are  well  defined  and  each  bears  a  single 


134  VERRILL 

spine  at  first.  There  are  no  actinal  spines  in  the  smaller  of  these 
(18  mm.  to  20  mm.  in  diameter),  but  one  row  appears  in  those 
about  23  mm.  to  25  mm.  in  diameter,  and  in  these  the  upper  mar- 
ginal plates  usually  begin  to  have  two  or  three  spines. 

This  species  has  been  very  much  confused  and  misunderstood  by 
writers,  owing  to  the  brief  and  poor  description  given  by  Brandt. 
He  described  many  of  his  North  Pacific  invertebrates  merely  from 
colored  drawings  made  by  or  for  Mertens,  and  probably  he  had  no 
specimens  in  most  of  such  cases.  The  starfishes  were  apparently 
among  those  thus  described  from  drawings  only,  for  several  were 
named  from  their  colors  and  in  some  cases  very  little  else  was  given. 
In  some  cases  Brandt  stated  that  the  specimens  were  lost.  His 
A.  ochracea  and  A.  epichlora  were  from  Sitka,  and  we  may  naturally 
conclude  that  they  would  be  species  commonly  found  there. 

About  A.  ochracea  there  can  be  no  doubt,  for  it  is  very  common 
on  the  rocks  at  Sitka  and  its  color  is  usually  characteristic,  as  well 
as  the  coarse  spinulation.  But  Dr.  Coe  informs  me  that  the  most 
abundant  species  between  tides  at  Sitka,  and  the  adjacent  coasts,  is 
the  one  now  under  discussion,  and  he  also  states  that  it  is  usually 
dull  green  or  olive-green  on  the  dorsal  surface  when  living,  which 
would  at  once  explain  the  name  epichlora.  Dr.  Coe  states  that  it 
has  the  same  habits  and  nearly  the  same  dull-green  color  as 
L.  littoralis  (St.),  which  he  had  seen  in  abundance  at  Eastport, 
Maine,  and  that  this  Alaskan  species  occurs  by  thousands  in  the 
same  way  at  Sitka,  on  the  rocks  at  low  tides.  It  also  carries  its  eggs 
and  young  in  clusters  over  the  mouth,  like  L.  littoralis.  These  facts 
would,  of  themselves,  make  it  almost  certain  that  Brandt's  species 
was  this  common  green  species,  especially  as  no  other  distinctly 
green  species  is  known  at  that  locality. 

Moreover,  Brandt's  description  fits  this  species  in  other  respects 
better  than  it  does  any  other.  The  principal  point  of  difference  is 
that  he  stated  that  it  was  five-rayed,  while  much  the  larger  propor- 
tion of  our  Sitka  specimens  are  six-rayed.  But  we  also  have  many 
five-rayed  specimens  of  exactly  the  same  character  from  Sitka.  As 
there  is  no  evidence  that  Brandt's  description  or  Mertens's  figure 
were  based  on  more  than  a  single  specimen,  this  difference  is  of  very 
little  importance.  Among  the  young  carried  by  six-rayed  mothers, 
are  found  more  or  less  five-rayed  ones. 

Brandt's  description  was  essentially  as  follows: 

"  The  disk  is  of  moderate  size  (about  one  inch  in  diameter),  sub- 
depressed.  The  five  arms  are  conical,  subdepressed,  unequal  in 


SHALLOW- WATER   STARFISHES  135 

length.  The  dorsal  surface  is  green,  covered  with  whitish  spines,  of 
which  the  summit  is  capitate,  arranged  crowdedly  in  a  reticulate 
manner.  Ventral  surface  pale  rose.  Diameter  of  disk,  one  inch; 
length  of  ray,  three  inches  [probably  measured  from  center  of 
disk]." 

Many  of  our  specimens  agree  very  well  with  this  in  size  and  pro- 
portion. But  the  young  of  troschelii,  when  of  this  size,  would  have 
a  very  small  disk  and  far  more  slender  and  longer  rays,  which  no 
one  could  call  "  conical "  or  "  subdepressed." l 

Several  writers  have  confounded  Brandt's  species  with  troschelii, 
especially  more  recently.  M.  de  Loriol  (1897)  has  given  a  full 
description  and  excellent  figures  of  troschelii,  under  the  name  of 
epichlora.  The  only  points  of  agreement  are  the  reticulate  arrange- 
ment of  the  dorsal  spines  and  their  capitate  form.  In  troschelii  the 
rays  are  not  only  much  longer  and  well  rounded  in  life,  but  its  color 
above  is  decidedly  red  or  brown.  Moreover,  although  it  occurs  at 
Sitka,  it  is  far  less  common. 

The  species  described  and  figured  by  De  Loriol  (op.  cit.,  p.  231, 
1897)  under  the  name  of  saanichensis,  from  Vancouver  Island, 
appears  to  be  the  five-rayed  variety  of  Brandt's  species,  agreeing 
very  closely  with  his  description.  It  also  agrees  very  closely  with 
some  of  our  Sitka  specimens. 

Brandt's  original  Latin  description  of  A.  camtschatica  (Prod., 
p.  270)  was  essentially  as  follows :  Diameter  of  disk  about  one  inch. 
Rays  six,  conical  acuminate,  scarcely  subequal  in  length,  one  to  one 
and  a  quarter  inches  long,  much  broader  toward  the  base.  The 
whole  back  dusky  or  ochraceous  and  covered  with  crowded,  pedi- 
cellate, truncate-capitate,  white  or  fuscus-white  spines,  arranged  in 
raised  reticulations,  forming  subregular  radial  series.  He  stated 
that  the  specimen  obtained  by  Mertens  had  been  lost,  but  had  been 
well  drawn  by  Postelesius,  and  that  there  were  other  specimens  in 
the  museum  of  the  Academy.  His  first  description,  however,  seems 
to  have  been  based  entirely  on  the  drawing.  Later  ( Middendorff's 
Reise,  1851,  u,  p.  32)  he  gave  a  rather  shorter  description  in  Ger- 
man, which  does  not  exactly  agree  with  the  former,  but  supplements 
it.  According  to  the  later  description  it  has  two  rows  of  ventral 
spines  and  three  of  adambulacrals  (probably  due  to  alternate  plates 
bearing  either  one  or  two  spines),  and  six  or  seven  rows  of  crowded 
dorsal  spines.  Greatest  diameter  of  the  largest,  three  inches; 
breadth  of  the  disk,  one  inch ;  length  of  arms,  one  inch  and  two  to 
four  lines. 

1  See  description  below,  page  155- 


136  VERRILL 

Both  of  his  descriptions  apply  very  well  indeed  to  many  of  the 
ordinary  six-rayed  specimens  of  L.  epichlora  alaskensis,  so  abun- 
dant on  the  coast  of  Alaska.  This  agreement  is  seen  not  only 
in  the  colors,  but  also  in  the  size  and  proportions  of  rays  to  the  disk, 
the  shape  and  the  crowded  reticulate  arrangement  of  the  spines,  etc., 
while  they  often  form  more  or  less  evident  rows,  as  in  var.  plena. 
This  arrangement  is  characteristic  of  many  specimens  of  epichlora, 
but  not  often  of  acervata  or  borealis.  Moreover,  the  longer  arms  and 
smaller  disk  of  the  latter  are  at  variance  with  the  proportions  given 
by  Brandt. 

Hence,  it  seems  very  probable  that  his  species  is  either  the  six- 
rayed  form  of  epichlora  or  else  some  very  similar  Siberian  species, 
and  therefore  quite  distinct  from  acervata  or  borealis.  It  might, 
possibly,  be  the  same  as  our  A.  multiclava.  The  description  is  insuffi- 
cient for  real  identification. 

LEPTASTERIAS  EPICHLORA  ALASKENSIS  Verrill,  nov. 

Plate  xvi,  figures  5,  6;  plate  xxvm,  figures  I,  2;  plate  LXXXV,  figures  i-id 
(details),  2-2e  (young). 

This  is  the  most  common  and  most  normal  form  of  the  species. 
It  differs  from  the  type  of  Brandt  mainly  in  being  six-rayed.  Disk 
of  moderate  size,  high,  swollen,  or  plump-looking.  Rays  six,  some- 
times five,  rather  short  and  stout,  rounded.  The  type  specimen 
has  the  radii  14  mm.  and  46  mm. ;  ratio,  about  1 :  1.33. 

Actinal  spines  in  two  or  three  rows,  terete  or  subclavate,  larger 
than  dorsals.  Dorsal  and  lateral  spines  short  and  very  numerous; 
the  dorsal  spines  unequal,  smaller  and  shorter  than  the  laterals, 
crowdedly  arranged  in  an  areolated  or  reticulated  pattern,  usually 
not  forming  definite  median  rows  except  when  young;  their  tips 
clavate  or  capitate,  spinulose  and  striated.  Large*  stout,  erect, 
wedge-shaped  dermal  major  pedicellariae  are  sparsely  scattered  over 
the  back  and  sides,  especially  in  the  lateral  grooves;  they  are  often 
as  broad  as  the  adjacent  spines  and  have  wide,  obtuse,  or  truncated, 
serrate  jaws.  (See  pi.  LXXXV,  figs,  i-irf.) 

The  madreporic  plate  is  usually  quite  small  and  inconspicuous, 
with  comparatively  few  coarse  gyri ;  but  in  some  of  the  larger  speci- 
mens, especially  five-rayed  ones,  it  is  much  larger,  with  numerous 
fine  gyri.  It  is  nearly  always  closely  surrounded  by  a  circular  group 
of  small  capitate  spines. 

There  is  a  complete  series  of  peractinal  ossicles,  each  bearing  one 
large  spine.  Large  specimens  may  have  a  rudimentary  row  of  small, 
usually  spinulose,  subactinals  near  the  base  of  the  rays. 


SHALLOW- WATER    STARFISHES  137 

Sitka,  Fox  Cape,  Yakutat,  Dutch  Harbor,  Aleutian  Islands  and 
Wrangel  (Harriman  Expedition).  Common  at  Sitka  at  low  tide. 
Many  of  the  specimens  have  large  clusters  of  very  young  ones 
beneath  them  over  and  around  the  mouth.  (PI.  LXXXV,  figs.  2-22.) 
Queen  Charlotte  Islands ;  Vancouver  Island ;  Puget  Sound ;  etc. 

Among  the  young  attached  under  the  mouth  of  the  mother,  were 
both  six-rayed  and  five-rayed  ones ;  the  former  much  more  numerous. 

Among  630  young,  taken  from  several  six-rayed  mothers,  and 
carefully  counted,  there  were  596  of  the  six- rayed  sort;  31  five-rayed ; 
3  four-rayed.  I  have  seen  no  adults  that  are  four-rayed.( 

A  six-rayed  specimen  from  St.  Paul's  Island  (coll.  H.  W.  Elliott, 
U.  S.  Nat.  Mus.,  No.  3267)  is  somewhat  peculiar.  It  is  90  mm.  in 
diameter,  with  the  rays  narrower,  more  depressed  and  more  acute 
than  usual,  so  that  it  resembles  A.  borealis  in  form.  But  the  dorsal 
spines  are  much  as  in  alaskensis,  very  numerous,  crowded,  arranged 
in  a  reticulate  or  areolate  manner,  and  all  capitate.  The  mar- 
ginal rows  are  double,  and  also  the  interactinal  row  proximally. 
These  spines  are  stout,  clavate.  The  adambulacral  spines  are  stouter 
than  usual,  especially  the  outer  ones,  and  decidedly  clavate.  On  the 
sides  of  the  rays  are  a  few  large,  erect,  blunt  pedicellariae,  nearly  as 
stout  as  the  adjacent  spines.  This,  like  several  other  odd  specimens, 
may  be  a  hybrid  between  L.  epichlora  and  A.  borealis. 

LEPTASTERIAS   EPICHLORA  ALASKENSIS,   Var.   CARI- 
NELLA  Verrill,  nov. 

Plate  xvi,  figures  I,  2. 

Normally  six-rayed.  Rays  short,  rounded.  Dorsal  ossicles  and 
spines  small,  capitate,  arranged  as  in  alaskensis,  except  that  there  is 
a  distinct  median  radial  row,  larger  than  the  rest.  Infero-marginal 
and  oral  spines  as  in  typical  specimens.  Large  erect  major  pedi- 
cellariae are  present  sometimes. 

Most  of  the  examples  of  this  variety  are  young,  up  to  50  mm.  in    fokt&krS 
diameter.    Probably  there  is  a  general  tendency  for  the  median  dorsal      ~ 
row  of  spines  to  become  indistinct  in  larger  specimens,  though  this  is 
not  always  the  case  in  this  species. 

Sitka  and  Dutch  Harbor,  Alaska  (Prof.  W.  R.  Coe,  Harriman 
Expedition). 

LEPTASTERIAS  EPICHLORA  ALASKENSIS,  Var. 
SIDEREA  Verrill,  nov. 

Plate  xvi,  figures  3,  4- 

Disk  large ;  rays  usually  six,  rather  short  and  stout.  Radii,  14  mm. 
and  45  mm.;  ratios,  about  1:3.2;  breadth  of  rays  at  base  12  mm. 

ft 


138  VERRILL 

Dorsal  spines  very  numerous  and  so  crowded  in  the  rows  and 
groups  that  their  capitate  tips  are  nearly  in  contact.  The  larger  ones 
are  nearly  even  in  size  and  height  over  the  whole  dorsal  and  lateral 
surfaces ;  they  are  short,  stout,  strongly  capitate,  with  roundish  finely 
spinulose  tips.  Along  the  median  lines  they  form  radial  ranges  or 
three  or  four  irregular,  crowded  rows ;  between  these  and  the  usually 
double  marginal  rows,  they  are  irregularly  reticulated,  or  stand  in 
little  groups  and  short  oblique  rows.  Small  ones  of  the  same  form 
are  scattered  between  the  larger  ones.  Ventral  spines  are  longer, 
rather  stout,  clavate,  arranged  mostly  in  three  rows,  two  of  which 
are  inferomarginals  and  one  peractinal. 

Adambulacral  spines,  slender,  slightly  clavate  or  obtuse,  either 
one  or  two  to  a  plate  irregularly.  They,  bear  groups  of  minor  pedi- 
cellariae. 

Dermal  major  pedicellariae  very  few,  mostly  marginal  and  inter- 
actinal,  small,  long-ovate;  smaller  acute-ovate  ones  are  attached 
within  the  ambulacral  grooves.  None  of  the  large,  wedge-shaped, 
serrate  ones  were  found  on  any  part. 

The  dorsal  skeleton  is  firm,  composed  of  larger  and  smaller  imbri- 
cated ossicles.  Papulae  rather  numerous  and  small  for  a  Leptasterias. 

In  form  and  proportions  this  is  very  similar  to  alaskensis,  but 
it  is  so  different  in  the  size  and  arrangement  of  its  dorsal  spines  that 
in  the  absence  of  a  series  of  specimens  I  should  have  supposed  that 
the  differences  of  spinulation  might  be  specific.  The  major  pedicel- 
lariae are  somewhat  different  in  the  two.  The  types  of  this  variety 
have  none  of  the  remarkably  large,  stout,  dentate  ones,  so  char- 
acteristic of  alaskensis.  A  large  series  of  specimens,  however, 
shows  that  this  is  only  a  peculiarity  of  certain  specimens. 

Sitka,  Dutch  Harbor,  and  jYakutatj  Alaska  (Dr.  W.  R.  Coe,  Har- 
riman  Expedition) ;  Puget  Sound. 

LEPTASTERIAS  EPICHLORA  MILIARIS  Verrill,  subsp.  nov. 

This  is  a  small  variety  with  pretty  numerous,  small,  crowded, 
slender  spinules,  so  that  it  resembles  L.  cequalis.  Rays  usually  six, 
sometimes  five.  Radii,  5  mm.  and  16  mm. ;  ratio,  about  1 :  3.2.  The 
median  radial  ossicles  bear  small,  slightly  larger,  capitate  spines 
surrounded  by  a  group  of  smaller  ones.  Other  dorsal  ossicles  usually 
bear  a  single  small,  capitate  spine  with  a  cluster  of  minute  ones 
i  around  it.  The  superomarginal  plates  bear  one  larger  spine  and  a 

number  of  unequal  smaller  ones,  several  of  which  stand  on  the 
descending  apophysis.    The  inferomarginals  usually  bear  two  small 


SHALLOW-WATER   STARFISHES  139 

divergent  spines.  The  peroral  and  adoral  spines  are  all  small  and 
slender,  as  in  the  young  of  the  more  typical  forms. 

Gape  Fox  and  Sitka  (Harriman  Expedition) ;  Queen  Charlotte 
Islands  (Canadian  Geological  Survey). 

This  variety,  which  is  remarkable  for  its  small  and  uniformly 
crowded  spinules,  is  liable  to  be  mistaken  for  L.  aqualis,  which  it 
closely  resembles  dorsally.  It  can  easily  be  distinguished  by  its 
much  feebler  oral  spines,  which  in  the  latter  are  larger  and  stronger 
than  usual  in  this  group. 

Variety  REGULARIS  Verrill,  nov. 

Rays  usually  six,  longer  and  more  slender  than  usual,  regularly 
rounded  and  gradually  tapered.  Radii,  6  mm.  and  25  mm.  Dorsal 
spines  numerous,  but  not  crowded,  nearly  all  capitate,  and  less 
unequal  in  size  than  in  most  varieties.  The  dorsal  radial  row  is  *"  7 
distinct,  each  ossicle  bearing  two  to  four  spines ;  two  or  three  stand 
together  on  many  other  ossicles.  Both  marginal  rows  and  one  inter- 
actinal  row  mostly  simple,  one  spine  to  a  plate.  Adambulacral, 
adoral,  and  oral  spines  as  in  other  varieties.  The  second  short 
interactinal  row  of  ossicles,  bearing  spines,  is  not  present,  as  it  is 
usually  in  other  varieties  when  of  the  same  size,  and  this  is  prob- 
ably the  cause  of  the  slenderness  of  the  rays. 

Cape  Fox,  Alaska  (Harriman  Expedition). 

Variety  SUBREGULARIS  Verrill,  nov. 

The  type  is  a  small  six-rayed  specimen  from  Sitka,  Alaska  (Prof. 
W.  R.   Coe,  Harriman  Expedition).     It  has  the  same  form  as        *-/*' 
alaskensis.    Its  marginal  spines  are  different.    The  superomarginal  W^  &** 
row  is  regular  and  nearly  simple,  only  one  spine  usually  standing  on 
a  plate ;  these  spines  are  unusually  large  for  this  group,  stout,  capi- 
tate, well  spaced.     The  inferomarginals  are  similar,  but  a  little 
longer.    There  are  two  interactinal  rows. 

The  dorsal  spines  are  short,  thick,  mammilliform  or  capitate;  the 
larger  ones  form  a  distinct  median  row;  others,  similar,  are  scat- 
tered, and  not  numerous.  The  papular  areas  are  large.  The  adam- 
bulacral  spines  are  rather  stout,  subclavate,  irregularly  subdipla- 
canthid. 

Variety  SUBNODULOSA  Verrill,  nov. 

At  first  sight  this  variety  might  easily  be  mistaken  for  the  young 
of  Pisaster  ochraceus,  on  account  of  the  conspicuous  and  prominent     — 
clusters  of  capitate  spines  scattered  over  the  dorsal  surface,  but  they 
do  not  form  reticulations. 


14O  VERRILL 

•^•*V 

The  type  is  five-rayed  and  larger  than  is  usual  in  this  species.  It 
equals  the  large  six-rayed  specimens  of  the  other  varieties.  The 
radii  are  10  mm.  and  60  mm. ;  ratio,  i :  6,  for  the  longer  rays.  The 
rays  are  decidedly  unequal,  however,  in  the  type,  owing  to  former 
injury.  The  arrangement  of  the  dorsal  ossicles  seems  to  be  much  as 
in  the  typical  form,  and  alaskensis,  but  the  spines  are  very  un- 
equal. The  smaller  are  very  numerous,  rather  minute  and  capi- 
tate, while  the  larger,  clustered  ones  are  larger  than  usual,  with 
broadly  capitate  or  button-like  ends,  wider  than  high.  The  clusters 
are  most  numerous  along  the  middle  of  the  rays,  but  do  not  form  a 
regular  row;  elsewhere  they  are  very  irregularly  scattered.  The 
superomarginal  spines  form  a  pretty  regular  row  of  slightly  longer 
blunt  spines  on  some  of  the  rays,  but  are  quite  irregular  on  others. 

The  single  row  of  inferomarginals  and  two  rows  of  interactinals 
are  longer,  not  so  stout,  bent  upward  a  little,  and  blunt  at  the  tip. 
The  second  row  of  interactinals  may  reach  nearly  to  the  end  of  the 
rays.  The  adambulacral  spines  are  as  in  var.  alaskensis.  Papulae 
stand  between  all  these  rows  of  spines,  either  in  small  groups  or 
singly.  Minor  pedicellariae  are  abundant  on  the  adambulacral  spines. 
Major  pedicellariae  are  few  and  rather  small,  ovate-lanceolate,  acute. 
None  of  the  large,  erect,  serrate  kind  were  observed. 

The  madreporic  plate  is  larger  than  is  usual  in  this  species  and  has 
thinner  and  more  numerous  gyri,  probably  due  to  greater  age. 

When  superficially  examined  this  looks  like  a  distinct  species. 
Although  the  dorsal  spinulation  appears  so  different,  it  is  arranged 
nearly  on  the  same  plan  as  in  more  typical  varieties.  The  unusual 
enlargement  of  the  larger  spines  in  the  clusters  is  only  an  exagger- 
ation of  a  tendency  present  in  most  of  the  larger  typical  specimens, 
which  often,  also,  have  the  more  enlarged  and  finer  madreporic 
plate.  The  marginal  and  actinal  plates  are  nearly  as  in  some  more 
typical  specimens.  It  may  possibly  be  a  hybrid  between  epichlora 
and  troschelii. 

Wrangel,  Alaska,  July  6,  1899  (Harriman  Expedition),  type. 

LEPTASTERIAS  EPICHLORA  PLENA  Verrill,  subsp.  nov. 
Plate  LVIII.  figure  i ;  text-figure  6. 

Rays  six  in  the  type,  rather  stout  and  regularly  tapered.  Radii 
are  13  mm.  and  43  mm. ;  ratio,  i :  3.30. 

The  dorsal  spines  are  very  numerous,  areolate,  short,  subequal, 
capitate,  with  rough  tops,  arranged  so  as  to  show  five  radial  bands 
on  irregular  multiple  rows,  separated  by  large,  papular  areas,  in  six 


SHALLOW-WATER   STARFISHES 


141 


obvious  radial  rows,  three  on  each  side.  The  median  row  of  spines 
is  distinct,  but  not  prominent.  The  dorsal  ossicles  are  openly  reticu- 
late, several  surrounding  each  papular  area,  and  bearing  spines  which 
are  so  numerous  as  to  be  nearly  in  contact  in  circles  around  the 
areas.  They  are  not  very  diverse  in  size.  Six  to  eight  often  stand 
in  a  cluster  on  the  larger  ossicles. 


FIG.  6. 


Leptasterias  epichiora  plena,  type,  a.  Group  of  dorsal  spines;  b,  b',  a  pair  of  supero* 
marginal  spines  on  one  plate;  p.  pedicellariae;  c,  c' ,  inferomarginal  spines;  d,  peractina! 
e,  outer,  and  f,  inner  adambulacral  spines  with  (/>)  minor  pedicellariae.  x  24. 

The  superomarginal  spines  are  similar  and  stand  in  groups  of 
two  to  four  on  a  plate,  besides  one  on  the  descending  apophysis.  The 
two  rows  of  marginal  ossicles  are  separated  by  a  row  of  large 
papular  areas,  bearing  small,  acute  major  pedicellariae. 

The  inferomarginal  spines  are  longer,  stouter,  clavate,  mostly 
somewhat  flattened,  obtuse.  They  mostly  stand  obliquely,  two  to  a 
plate,  forming  a  crowded  double  row.  The  peractinal  spines  are 
stouter,  shorter,  more  clavate  and  more  flattened,  one  to  a  plate,  close 
to  the  adambulacrals.  The  latter  are  much  more  slender,  two  to  a 
plate.  The  outer  ones  are  a  little  shorter  than  the  inner,  stouter,  and 


2- 
Ib 


142  VERRILL 

mostly  a  little  clavate  and  flattened.  The  inner  one  is  slender, 
tapered,  and  often  bears  a  small  cluster  of  acute  pedicellariae. 
Adoral  spines  are  similar  but  longer.  No  large  dermal  major  pedi- 
cellariae were  found. 

Vancouver  Island  (Canadian  Geological  Survey). 

The  type  of  this  is  a  very  distinct  form.  The  equal,  crowded, 
capitate  dorsal  spines,  arranged  in  areolations  and  evident  longi- 
tudinal bands,  are  notable;  the  superomarginals,  three  or  four  to  a 
plate,  are  also  peculiar.  The  open  reticulation  of  the  ossicles  is  per- 
haps more  important. 

LEPTASTERIAS  EPICHLORA  PUGETANA  Verrill,  subsp.  nov. 

The  type  specimen  from  Puget  Sound  is  very  peculiar  in  appear- 
ance. The  dorsal  spines  are  nearly  equal,  numerous,  crowded,  and 
capitate.  They  are  somewhat  areolate,  forming  circles  around  the 
very  evident  papular  areas.  The  median  radial  bands  are  evident, 
but  not  prominent.  Both  the  marginal  rows  are  double,  with  rather 
stout  obtuse  spines.  The  peractinal  row  has  similar  short  spines, 
close  to  the  adambulacrals.  The  skeletal  ossicles  are  stout  and 
rather  closely  articulated.  Large,  erect,  serrate  major  pedicellariae 
are  numerous  on  the  sides  of  the  rays.  Radii,  10  mm.  and  35  mm. ; 
ratio,  i :  3.5. 

Puget  Sound  (Professor  Ritter,  Museum  of  the  University  of 
California). 

LEPTASTERIAS  ( ?)  DISPAR  Verrill,  sp.  nov. 
Plate  xvi,  figure  7. 

Disk  moderately  large,  rays  six,  angular,  depressed,  rather  short, 
tapered  rapidly.  Radii,  8  mm.  and  32  mm. ;  ratio,  1 : 4.  Dorsal 
spines  very  unequal  in  size  and  form.  The  larger  ones  are  stout 
and  strongly  capitate,  with  large,  round,  finely  spinulose  heads ;  they 
form  conspicuous  but  irregular  median  radial  rows,  two  or  three 
often  standing  on  one  plate,  and  also  small  groups  on  the  disk. 
Smaller  spines  of  various  sizes,  partly  capitate  like  the  larger  ones, 
and  partly  slender  and  tapered  or  slightly  clavate,  are  scattered  over 
the  surface,  but  mostly  in  imperfect  submedian  rows;  larger  speci- 
mens might  have  them  in  regular  rows. 

Marginal  spines  are  similar  to  the  larger  dorsals,  but  not  so  stout. 
They  form  pretty  regular  rows,  one  to  a  plate  proximally,  but  mostly 
two  to  a  plate  distally.  Ventral  spines  are  decidedly  longer  and 


SHALLOW- WATER   STARFISHES  143 

more  slender,  tapered,  subacute;  they  stand  in  three  rows,  or  some- 
times four,  at  the  bases  of  the  rays.  Of  these  the  inferomarginals 
stand  mostly  two  to  a  plate  or  three  to  a  plate  proximally.  Per- 
actinal  spines  form  a  simple  row,  not  reaching  the  tip  of  the  ray. 
Adambulacral  spines  small,  about  half  as  long  as  the  inferomarginals, 
very  slender,  terete,  or  slightly  clavate,  mostly  two  to  a  plate ;  some- 
times alternately  one  and  two. 

Marginal  and  dorsal  dermal  major  pedicellariae  unusually  small, 
few,  compressed,  ovate  and  acute-lanceolate.  Many  small  ones 
occur  along  the  inner  edges  of  the  adambulacral  grooves.  Minor 
pedicellariae  are  rather  few  on  the  dorsal  spines,  but  more  numerous 
on  the  marginal  and  peractinal  spines ;  a  few  are  scattered  between 
the  spines  on  the  dorsal  surface. 

The  ambulacral  grooves  are  rather  shallow  and  wide,  appearing 
much  more  open  than  is  usual  in  other  species  of  this  size.  The 
pores  are  rather  large  and  not  much  crowded. 

The  mouth  is  not  sunken.  The  peroral  spines  are  well  developed, 
but  rather  short  and  divergent.  The  apical  pairs  are  rather  longer 
and  stouter  than  the  adambulacral,  tapered,  subacute;  those  of  the 
exterior  pair  are  of  the  same  form,  from  one-half  to  two-thirds  as 
long,  and  very  divergent.  The  epioral  and  adoral  pairs  are  dis- 
tinctly longer  than  those  further  out.  Three  contingent  pairs  of 
plates,  besides  the  epiorals,  form  the  adoral  carina.  The  oral  spines 
often  bear  small  oblong-lanceolate  major  pedicellariae. 

The  dorsal  ossicles  are  unusually  large  and  thick  for  so  small  a 
species.  They  are  wide  and  deeply  lobed  and  are  closely  and  firmly 
united  together,  leaving  only  very  small  spaces  for  the  papulae,  which 
are  few  and  often  stand  singly.  The  larger  plates  form  three  longi- 
tudinal rows  besides  the  superomarginals,  which  are  four-lobed,  con- 
spicuous, and  elevated  in  the  middle,  like  the  median  radials.  The 
latter  are  strongly  imbricated,  and  have  large  lateral  lobes. 

The  type  is  from  Dutch  Harbor,  Unalaska  (Harriman  Expedi-     rut_£ 
tion) . 

This  most  resembles  L.  inequalis,  except  in  having  six  rays.  The 
comparatively  large  and  thick  dorsal  ossicles,  angular  rays,  open 
grooves,  and  general  appearance  of  the  type  indicate  that  it  may  be 
the  young  of  a  species  that  grows  to  a  much  larger  size,  but  I  know 
of  no  species  to  which  it  could  be  referred,  unless  it  be  another  of  the 
numerous  varieties  of  aceruata  or  epichlora.  But  it  differs  widely 
from  all  those  forms  of  the  latter,  that  I  have  seen,  in  the  large, 
lobate,  dorsal  ossicles,  arranged  pretty  clearly  in  three  longitudinal 


144  VERRILL 

rows,  and  without  the  reticulations,  generally  characteristic  of 
epichlora.  The  inferomarginal  and  actinal  spines  are  much  longer 
than  in  the  latter.  The  ambulacral  grooves  and  pores,  the  oral 
spines,  and  the  adambulacral  spines  are  also  different,  so  that  the 
under  surface,  as  well  as  the  dorsal,  differs  decidedly  in  appearance 
from  any  of  the  recognized  forms  of  epichlora,  all  of  which  are 
much  alike  beneath,  however  much  the  dorsal  spines  may  vary.  The 
pedicellariae  also  appear  to  be  peculiar. 

It  more  nearly  resembles  some  of  the  undoubted  young  of 
A.  acervata,  but  the  lateral  and  ventral  spines  are  all  longer  and  more 
slender.  It  may  not  be  a  Leptasterias.  Genital  pores  not  observed. 

LEPTASTERIAS  OBTECTA  Verrill,  sp.  nov. 

Rays  five,  rather  short  and  rapidly  tapered  to  slender  tips.  Radii, 
6  mm.  and  25  mm. ;  ratio,  1 : 4.25. 

Whole  surface  closely  covered  with  minute,  nearly  equal,  short, 
rough-tipped  spinules,  surrounded  with  large,  dense  wreaths  of 
unusually  large,  blunt-ovate  minor  pedicellariae,  so  abundant  as  to 
cover  the  spaces  between  the  spines  and  largely  conceal  the  spines, 
except  the  tips,  thus  giving  the  surface  a  smoothish  appearance  like 
Henricia. 

The  dorsal  ossicles  are  very  numerous,  small,  irregularly  reticu- 
lated, leaving  many  small,  irregular  papular  areas ;  the  lateral  supra- 
marginal  areas  are  not  tansversely  elongated,  as  in  C.  cribraria.  The 
dorsal  spinules  stand  in  small  irregular  clusters  of  three  to  six  or 
more;  the  ossicles  are  without  much  tendency  to  form  transverse 
rows.  Carinals  not  easily  distinguished. 

Marginal  plates  small  and  quite  concealed  by  the  abundant  pedicel- 
lariae; those  in  each  row  usually  bear  two  small  spines,  sometimes 
three,  a  little  longer  than  the  dorsals.  Proximally  there  is  a  short 
row  of  intermarginals  and  of  interactinals,  each  bearing  one  small 
spine,  partly  covered  with  pedicellariae.  Adambulacrals  dipla- 
canthid,  the  spines  small,  slender,  slightly  clavate;  adorals  longer. 
The  minor  pedicellariae  have  bases  about  as  broad  as  the  spines ;  the 
blades  are  spatulate  and  rounded  at  the  tips.  The  dorsal  spinules 
are  mostly  four  to  six  times  as  long  as  broad ;  tips  are  microscopic- 
ally thorny.  Genital  pores  and  reproduction  unknown. 

Off  Kings  Island,  Bering  Sea,  in  seventeen  fathoms.  (Coll.  Mus. 
Comp.  Zool.,  No.  1208;  one  dry.) 


SHALLOW-WATER   STARFISHES  145 

Genus  Stenasterias  Verrill,  nov. 
Type,  5*.  macropora  Verrill. 

Rays  slender,  six  in  the  type.  Dorsal  plates  are  wide,  lobed,  imbri- 
cated, stichasterial  in  arrangement.  Median  radial  row  and  dorso- 
lateral  rows  are  nearly  regular. 

Both  marginal  rows  are  formed  of  wide  imbricated  plates.  One 
row  of  small  peractinal  plates,  and  a  short  subactinal  row.  All  these 
plates  are  closely  covered  with  clusters  of  minute  spinules.  Papular 
areas  small;  papulae  few,  often  isolated.  Adambulacral  and  ambu- 
lacral  plates  not  strongly  compressed.  Ambulacral  pores  unusually 
large,  in  four  rows ;  podia  large.  Adambulacral  plates  are  dipla- 
canthid.  Reproduction  unknown. 

STENASTERIAS  MACROPORA  Verrill.  ^ 

Plate  L,  figure  7;  plate  LXXIV,  figure  4  (type);  plate  LXXxW,  figures  5-5*  jU^ 

(details). 

Leptasterias  macropora  VERRILL,  Amer.  Journ.  Sci.,  xxvm,  p.  65,  fig.  10,  1909. 

Disk  small.  Rays  six,  long,  slender,  constricted  at  base,  convex, 
with  a  median  dorsal  row  of  more  prominent  ossicles.  Radii  are 
3  mm.  and  15  mm. ;  ratio,  1:5.)  Ambulacral  feet  and  pores  unus- 
ually large;  the  pores  are  triangular,  overlapping  by  their  acute 
angles,  and  separated  only  by  thin  plates.  Adambulacral  plates  are 
strong,  unusually  thick  radially.  Adambulacral  spines  mostly  two  to 
a  plate,  long,  slender,  tapered,  acute  or  subacute.  ( The  grooves  are 
unusually  wide  and  open.  A  single  row  of  somewhat  quadrate,  over- 
lapping interactinal  plates,  which  bear  one  or  two  small,  rather  stout, 
tapered  spines ;  a  series  of  small,  oblong  connective  ossicles  between 
these  and  the  adambulacral  plates,  on  the  basal  part  of  the  ray,  each 
of  which  may  also  bear  a  spine.  A  row  of  larger,  rhombic,  imbri- 
cated marginal  plates,  in  contact  with  or  overlapping  the  sides  of  the 
peractinals,  runs  along  the  under  side  of  the  rays,  but  curves  upward 
to  the  dorsal  side  at  the  disk.  These  lateral  plates  usually  bear  two 
or  three  small  tapered  spines,  like  the  actinals ;  between  them  and  the 
actinal  plates,  and  between  the  latter  and  the  adambulacrals,  there 
are  only  very  small  spaces,  usually  occupied  by  a  single  papula. 

The  dorsal  plates  are  also  rather  closely  imbricated,  the  spaces 
between  being  very  few  and  small,  with  only  one  or  two  papulae ;  the 
median  radial  plates  are  thicker  and  larger,  subtriangular,  with  acute 
cusps,  and  concave  edges.  These  overlap,  or  are  imbricated  upon, 
the  plates  proximal  to  them,  so  as  to  form  a  median  radial  ridge 
or  carina.  Other  dorsal  plates  are  smaller  and  more  irregular.  The 
ii 


146  VERRILL 

dorsal  spines  are  numerous,  small,  short,  stumpy,  usually  truncate  or 
slightly  clavate,  but  not  much  enlarged  at  tip  and  not  much  longer 
than  thick.  They 'are  in  small  groups  on  the  larger  plates,  isolated 
on  the  smaller  ones.  Major  pedicellariae  of  rather  large  size,  but 
few  in  number,  are  present  on  the  inferior  interbrachial  areas ;  these 
are  compressed,  acute,  lanceolate.  Others  of  similar  form,  but 
smaller,  are  found  within  the  edges  of  the  ambulacral  grooves  and 
on  the  adjacent  spines.  Minor  pedicellariae  are  few  and  minute. 

The  specimens  of  this  species  are  small  and  poorly  preserved, 
having  lost  many  of  their  spines.  It  appears  to  be  allied  to 
L.  cequalis  more  nearly  than  to  other  species,  but  it  is  much  more 
slender  than  that  and  has  much  larger  ambulacral  pores,  while  the 
dorsal  plates  are  closely  imbricated. 

Sitka  (Harriman  Expedition)  ;  Queen  Charlotte  Islands  (G.  M. 
Dawson) . 

Genus  Stephanasterias  Verrill. 
Type,  5".  albula  Verrill  (Stimpson  sp.). 

Stichaster  (pars)  VERRILL,  op.  cit,  1866,  p.  551 ;  Perrier,  op.  cit,  1875,  p.  347 ; 

Sladen,  op.  cit.,  1889,  p.  432. 
Stephanasterias  VERRILL,  Bull.  Essex  Inst.,  i,  p.  5,  1871 ;  op.  cit.,  1874,  p.  353 ; 

Revision  Genera,  op.  cit.,  1899,  p.  222. 
Nanaster  PERRIER,  op.  cit.,  1894,  pp.  129,  131,  133. 

Dorsal  ossicles  small,  not  closely  joined,  the  radial  series  not  in 
very  regular  rows;  median  row  more  or  less  imbricated;  often  not 
much  differentiated,  three-lobed;  transverse  ossicles  strong,  notably 
regularly  arranged  in  oblique  rows;  all  covered  with  numerous 
crowded,  small,  subequal  spinules,  in  divergent  clusters,  arranged  in 
transverse  rows  on  the  rays.  Papular  areas  serial,  small,  with  few 
or  solitary  papulae. 

Superomarginal  plates  similar  to  dorsals;  not  very  distinct,  and 
with  similar  numerous  spinules.  Inferomarginal  plates  oblique  and 
prominent,  in  a  close  row,  multispinose,  their  spines  longer  than  those 
of  dorsals.  No  interactinal  plates.  Adambulacral  spines  usually  dip- 
lacanthid  or  irregularly  triplacanthid.  Sucker-feet  quadriserial. 
Pedicellariae  of  two  kinds,  as  in  Asterias;  many  dermal  minor  pedi- 
cellariae. 

Besides  the  type,  which  is  autotomous,  this  genus  appears  to 
include  5\  gracilis  of  the  West  Indies,  which  is  six-rayed. 

These  two  species  have  usually  been  referred  to  Stichasterinae  by 
recent  writers,  but  to  me  they  seem  closely  allied  to  Leptasterias. 
(See  also  pp.  40-41,  above.)  Seen  from  the  inner  surface,  the  dorsal 


SHALLOW-WATER   STARFISHES  147 

ossicles  are  reticulated  rather  than  in  longitudinal  rows,  and  the 
transverse  ossicles  are  larger  than  the  radial ;  the  median  row  is  only 
a  little  larger  than  the  others.  The  papular  areas  are  wider  than  the 
plates.  Reproduction  unknown. 

STEPHANASTERIAS  ALBULA  (Stimpson)  Verrill. 
Asteracanthion  albulus  STIMPSON,  Invert.  Grand  Manan,  p.  14,  pi.  i,  fig.  5, 

^      1853- 

Stichaster  albulus  VERRILL,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  x,  p.  351,  1866; 
op.  cit,   1895,  p.  206.     Perrier,  Arch.  Zool.  Exper.,  vol.  iv,  p.  347,   1875. 

Duncan  and  Sladen,  op.  cit.,  p.  29,  pi.  2,  figs.  13-17,  1881.    Danielssen  and 

Keren,  op.  cit.,  p.  31,  pi.  8,  figs.  13-15,  1884.     Sladen,  Voy.  Challenger, 

vol.  xxx,  p.  432,  1889. 
Asteracanthion  problema  STEENSTRUP,  Vidensk.  Medd.  nat.  Foren.,  p.  240, 

1854.    Liitken,  Gronl.  Echinod.,  p.  30,  1857. 
Stephanasterias  albula  VERRILL,  Bulletin  Essex  Inst.,  vol.  i.  p.  5,  1871;  Expl. 

Casco  Bay,  p.  353,  1874;  Check  List,  1879;  Expl.  by  the  Albatross  in  1883, 

p.  540,  1885;  Revision  Genera  and  Species  Starfish,  Trans.  Conn.  Acad., 

vol.  x,  p.  222,  1899. 
Variety  nitida  VERRILL,  op.  cit.,  1866,  p.  351. 

A  small  autotomous,  finely  spinulated  species,  with  five  to  nine  or 
more  usually  unequal  rays,  normally  with  six  when  full  grown  (var. 
nitida) . 

In  the  small  specimens,  while  dividing,  there  are  usually  two  or 
three  longer,  and  two  to  four  shorter  rays.  Mature  specimens  may 
have  the  larger  radius  50  mm. ;  lesser,  6  mm. 

The  dorsal  plates  in  the  larger  specimens  bear  about  eight  to 
twelve  small,  crowded,  divergent,  costellate  and  rough-tipped  spin- 
ules ;  the  median  series  may  be  a  little  larger  and  distinct.  About  five 
close-set,  alternating  dorso-lateral  rows  of  clusters  of  spinules  proxi- 
mally;  inferomarginal  plates  bear  three  to  five  distinctly  larger  and 
longer  spines. 

Adambulacral  plates  bear  two  or  three  (sometimes  four  in  large 
specimens),  very  slender,  obtuse  spines. 

Major  pedicellarise  small,  lanceolate  or  ovate ;  they  occur  in  a  row 
on  the  inner  edge  of  the  ambulacral  furrow ;  on  the  interradial  areas ; 
and  sometimes  on  the  adambulacral  spines;  a  few  dorsal  ones  are 
much  larger,  stout-ovate,  twice  as  thick  as  the  spinules.  Minor  pedi- 
cellariae  small,  with  abruptly  spatulate  valves,  nearly  as  thick  as  the 
spinules,  numerous  between  the  spines,  above  and  below.  Papulae 
rather  large,  mostly  in  pairs  or  single. 

This  is  a  circumpolar  species.  Its  range  extends  to  Greenland, 
Iceland  and  other  parts  of  the  Arctic  Ocean,  and  the  northern  coasts 
of  Europe  and  Asia. 


148  VERRILL 

On  the  east  American  coast,  it  is  common  from  low-water  mark 
to  loo  fathoms  in  the  Bay  of  Fundy  and  off  the  coast  of  Nova  Scotia. 
Dredged  at  more  than  100  stations  between  N.  lat.  46°  50'  and 
35°  12'  30".  Off  Cape  Hatteras  and  off  S.  Carolina  it  is  common 
in  1 6  to  50  fathoms.  Common  south  of  Martha's  Vineyard  in  50  to 
150  fathoms. 

In  depth,  its  range  is  o  to  229  fathoms ;  in  one  case  recorded  from 
435  fathoms,  off  Delaware,  and  once  from  1253  fathoms. 

Prof.  W.  K.  Fisher  informs  me  that  he  has  North  Pacific  speci- 
mens from  the  following  localities :  Stephens  Passage,  Alaska,  Alba- 
tross Station  4253,  in  131  to  188  fathoms,  rocks  and  shells;  Shelikoff 
Strait,  Station  4292,  94  to  102  fathoms,  blue  mud  and  sand.  Off 
Bering  Island,  Station  4791,  in  76  fathoms,  rocks. 

The  specimen  examined  by  me  is  from  the  last  named  locality,  sent 
by  Professor  Fisher. 

A  closely  related  form,  S.  gracilis  (Per.),  occurs  in  deep  water  in 
the  West  Indies. 

Genus  Ctenasterias  Verrill,  nov.     (See  p.  53.) 
Type,  C.  spitzbergensis  (Danielssen  and  Koren,  sp.). 

CTENASTERIAS  CRIBRARIA  (Stimpson). 
Plate  xxv,  figures  3,  4  (type)  ;  text-figure  7. 

Asterias  cribraria  STIMPSON,  Proc.  Boston  Soc.  Nat  Hist,  vm,  p.  270,  1862. 

Ludwig,  Zool.  Jahr.,  p.  288,   1886;  Fauna  Arctica,  p.  482,  1900.     Bell, 

Asteriae,  1881,  pp.  494,  505. 
Asterias  spitsbergensis  DAN.  and  KOREN,  Norwegian  N.  Atlantic  Expedition. 

Asteroidea,  p.  5,  pi.  i,  figs,  i-n,  1884. 

This  small,  five-rayed  species  was  thus  described  by  Stimpson : 
"  This  species  has  very  much  the  aspect  of  a  Cribrella.  Body 
thick  and  tumid,  with  a  smoothish  appearance  resulting  from  the 
great  number  and  small  size  of  the  spines.  Skin  rather  pliable  or 
coriaceous ;  ossicles  very  slender,  but  very  numerous.  Rays  five,  not 
contracted  at  base,  and  rather  higher  than  broad.  Disc  rather  large. 
Proportion  of  the  diameters  i :  4.8.  Ambulacral  pores  not  crowded, 
and  forming  two  zigzag  rows  rather  than  four  distinct  rows.  Some 
small,  acutely  triangular  major  pedicellariae  in  the  furrows.  Ambu- 
lacral spines  cylindrical  with  bluntly  rounded  tips,  forming  (except 
toward  the  extremity  of  the  ray)  two  regular  rows,  two  to  each 
plate,  and  bearing  small  clusters  of  minor  pedicellariae  at  their  outer 
bases.  Ventral  and  lateral  spines  in  regular  rows,  but  passing  imper- 


SHALLOW-WATER    STARFISHES 


149 


ceptibly  into  the  dorsal  spines  on  the  high,  rounded  side  of  the  ray ; — 
four  of  these  rows  may  be  counted,  in  which  the  spines  are  small, 
slender,  shorter,  and  more  pointed  than  the  ambulacral  spines,  and 
surrounded  at  base  by  thick  wreaths  of  minor  pedicellariae,  which 
wreaths,  in  alcoholic  specimens,  touch  each  other  at  their  bases.  The 
dorsal  ossicles,  with  their  interspaces,  are  mostly  transverse  in  direc- 
tion on  the  rays,  and  anastomose  pretty  closely,  except  that  there  is 


X24 


FIG.  7. 


Ctenasterias  cribraria.  No.  6123,  U.  S.  Nat.  Mus.  (Young).  A,  A  jaw;  a,  a;  a',  a',  apical 
spines;  p,  p,  pedicellariae;  e,  e,  epiorals.  B,  Ventral  spines;  a,  a,  inner,  and  a'  a',  outer 
adambulacrals;  b,  peractinals;  c,  c,  inferomarginals;  p,  p',  minor  pedicellariae;  p",  adambu- 
lacral  pedicellariae;  X  24.  C,  A  group  of  dorsal  spines,  minor  pedicellariae,  and  papular 
pores,  x  24.  D,  Pedicellarias;  a,  two  of  the  minor  sort;  b,  c,  c',  small  major  pedicellariae. 
X  67. 

on  each  side  a  series  of  transverse  membranous  interspaces  much 
larger  than  the  rest  (often  one-fifth  the  width  of  the  ray)  and  each 
containing  from  two  to  five  papulae.  The  papulae  elsewhere  stand 
singly,  sometimes  two  together.  The  dorsal  spines  are  very 
numerous,  minute,  no  thicker,  and  much  shorter  than  the  latero- 
ventrals,  and  are  more  or  less  capitate ; — they  are  somewhat  variable 
in  size,  and  are  arranged  in  groups  on  the  ossicles.  Among  them  are 
considerable  numbers  of  minor  pedicellariae,  which  are  often  half  as 


I5O  VERRILL 

large  as  the  spines  themselves.  On  the  disc  the  spines  are  very 
much  crowded,  as  they  also  are  along  the  middle  of  the  ray,  forming 
a  more  or  less  distinct  median  series.  The  spines  of  the  eyelids  and 
extremities  of  the  rays  are  much  stouter  than  any  of  the  others  either 
above  or  below.  The  madreporic  plate  is  large,  but  not  surrounded 
by  any  special  arrangement  of  protecting  spines.  The  minor  pedi- 
cellariae  in  this  species  are  strongly  truncated  at  the  extremity.  The 
major  pedicellariae  are  few  in  number,  and  situated  on  the  disc 
below, — small  ones  at  the  inner  bases  of  the  labial  spines, — and  two 
or  three  very  large  ones  in  the  angle  of  the  rays ;  the  latter  having 
stout,  almost  cylindrical  valves,  one  of  which  is  sometimes  notched 
at  the  extremity  for  the  reception  of  the  point  of  the  other.  A  large 
specimen,  probably  of  this  species,  was  found,  in  which  some  of  these 
large  major  pedicellarise  also  occurred  on  the  sides  of  the  rays. 
Diameter,  usually  two  and  one-half  inches. 

"  This  fine  species  appears  to  be  allied  to  A.  Miilleri,  although  so 
different  in  aspect. 

"  Dredged  in  considerable  numbers  on  a  muddy  bottom  in  from 
twenty  to  thirty  fathoms,  in  the  Arctic  Ocean,  north  of  Bering's 
Straits.  U.  S.  North  Pacific  Expedition.  Capt.  John  Rodgers." 

St.  Matthew's  Island;  Lawrence  Bay,  in  15  to  20  fathoms 
i*.  &/•  (Ludwig). 

Natural-size  photographs,  furnished  by  Dr.  Rathbun,  are  here 
reproduced.  They  are  from  Stimpson's  type,  in  the  U.  S.  National 
Museum. 

Ludwig  (op.  cit),  1886,  gives  the  following  information: 

"  The  present  examples  of  this  species,  known  hitherto  only 
northerly  from  Bering  Strait,  from  a  depth  of  20  to  30  fathoms,  were 
discovered  southerly  from  Bering  Strait.  The  largest  specimen 
(6.5  cm.  across)  comes  from  St.  Matthew's  Island;  the  remaining 
seven  specimens  are  from  Lorenz  Bay.  Among  the  latter  is  found 
one  5  cm.  across,  which  possesses  only  four  arms,  while  all  the 
others  are  five-armed.  With  the  exception  of  the  four-armed  indi- 
vidual, the  specimens  from  Lorenz  Bay  are  young  animals,  which 
have  a  size  of  from  2  cm.  to  3.3  cm.  In  five  cases  the  depth  and  con- 
dition of  the  bottom  are  given;  the  former  amounts  to  15  to  17 
fathoms ;  the  latter  is  designated  as  '  fine  clay  mud.'  " 

This  appears  to  be  a  strictly  arctic  and  circumpolar  species.  It  has 
not  been  taken  on  the  southern  coast  of  Alaska,  so  far  as  I  know. 

The  Asterias  spitzbergensis  Dan.  and  Koren  must  be  very  closely 
related  to  this,  and  probably  identical.  It  has  the  same  general 


SHALLOW- WATER   STARFISHES  15 1 

appearance,  due  to  the  large  number  of  small,  even  spinules,  and 
their  peculiar  transverse  arrangement  on  the  sides  of  the  rays;  the 
same  reticulate  arrangement  of  slender  dorsal  ossicles ;  and  essen- 
tially the  same  arrangement  of  the  marginal  ossicles  and  spines. 
Superomarginal  ossicles  seem  to  be  equally  obscured  in  both.  Both 
also  have  three  corresponding  forms  of  pedicellariae.  In  both, 
the  forcipulate  ones  are  unusually  stout  and  blunt ;  in  both  there  are 
adambulacral  forficulate  pedicellariae  with  acute  tips,  and  also  a 
much  larger  dermal  kind,  "  nearly  cylindrical,"  with  wide  obtuse 
valves.  The  last-named  form  is  very  unusual  and  is  a  strong  indi- 
cation of  specific  identity.  The  principal  difference  seems  to  be  that 
in  C.  cribraria  the  adambulacral  plates  are  described  as  having  two 
spines,  while  in  C.  spitzbergensis  they  bear  alternately  two  and  three 
spines.  The  differences  may  well  be  due  to  the  much  larger  size 
of  the  latter,  which  was  about  twice  as  large  as  the  former. 

After  carefully  comparing  specimens  of  both,  my  conclusion  is  that 
they  are  identical  but  variable. 

Several  young  specimens  of  cribraria  of  different  sizes  were  sent 
to  me  by  the  United  States  National  Museum  (No.  6123).  These 
were  taken  in  the  Arctic  Ocean,  north  of  Alaska,  near  Icy  Cape,  in 
10  to  15  fathoms,  mud  and  sand.  Smith,  1874;  coll.  W.  H.  Dall,  No. 
1 212.  One  of  these  furnished  the  details  shown  in  text-figure, 
No.  7.  The  larger  of  these  (greater  radius,  30  mm.)  agrees  almost 
exactly  with  a  specimen  of  similar  size  from  Spitzbergen,  17  to  23 
fathoms,  July  25,  1899,  sent  by  the  Museum  of  Comparative  Zoology. 
Another,  from  Greenland,  sent  with  last,  is  smaller  and  agrees  well 
with  the  smaller  Alaskan  specimens.  These  were  labelled  as  A. 
grcenlandica,  but  do  not  agree  with  typical  specimens  of  the  latter 
sent  to  me  by  Dr.  Chr.  Lutken. 

Genus  Evasterias  Verrill,  nov. 

A  group  of  long-rayed,  diplacanthid  starfishes  with  a  rather  small 
disk  and  a  reticulated  dorsal  skeleton,  bearing  numerous  small  spines 
arranged  in  a  reticulated  or  subreticulated  pattern,  or  in  transverse 
combs,  and  having  several  rows  of  imbricated  interactmal  ossicles,  of 
which,  in  the  adult,  there  may  be  three  to  five  or  more  rows.  Type, 
E.  troschelii  (Stimpson).  (See  also  p.  51.) 

EVASTERIAS  TROSCHELII  (Stimpson). 
Plate  xxu,  figures  i,  2;  plate  xxv,  figures  I,  2  (type)  ;  plate  xxvi,  figures  I,  2 

(typical)  ;  plate  LXII,  figure  i  (variety)  ;  plate  cvi,  figures  i,  2  (young). 
Asterias  troschelii  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  vin,  p.  267,  1862. 
Verrill,  Trans.  Conn.  Acad.,  n,  part  2,  p.  326  (no  descr.)  1868.    Perrier, 


152  VERRILL 

op.  cit.,  1875,  P-  335  (no  descr.).    Bell,  1881,  pp.  495,  505.    Whiteaves, 

Trans.  Royal  Soc.  Canada,  iv,  p.  116,  1887. 
Asterias  brachiata  Perrier,  Arch.  Zool.  Exper.,  iv,  pp.  329,  357,  1875  (locality, 

Gulf  of  Georgia)  (non  L.). 
Asterias  (Diplasterias)  epichlora  DE  LORIOL.  Mem.  Soc.  Phys.  et  Hist.  Nat., 

Geneve,  xxxii,  p.  19,  pi.  in,  figs.  2-2d,  187!  (non  Brandt). 

Several  specimens  which  evidently  belong  to  this  species,  were  col- 
lected by  the  Harriman  Expedition. 

When  well  grown  it  is  a  large,  five-rayed  starfish  with  a  small  disk 
and  long,  round,  tapering  rays,  with  a  rough  uneven  dorsal  surface, 
covered  with  short,  unequal,  obtusely  rounded  or  capitate  spines, 
which  are  arranged  in  an  irregularly  reticulated  or  areolated  pat- 
tern, and  always  in  short  rows  and  more  or  less  clustered  or  acer- 
vate.  The  inf eromarginal  and  actinal  plates  bear  usually  four  to  six 
or  more  rather  regular  rows  of  longer  and  larger,  mostly  blunt  spines. 
Two  irregular  series  of  slender  adambulacral  spines. 

Dr.  Rathbun  has  forwarded  two  photographs  (here  reproduced, 
pi.  xxv)  of  a  part  of  one  ray  of  the  type,  which  seems  to  be  all  that 
is  preserved  in  the  U.  S.  National  Museum.  Very  little  can  be  added 
to  the  original  description  from  this  fragment.  This  specimen  was 
less  than  half  the  full  size.  The  original  description  was  as  follows : 

"  Rays  five,  slender  and  somewhat  pentagonal,  regularly  tapering 
to  a  point ;  disc  small.  Proportion  of  the  diameters,  i :  7.  Ambu- 
lacral  pores  in  four  regular  rows.  Ambulacral  spines  in  two  or  three 
rows,  generally  two,  but  occasionally  one,  to  each  plate;  they  are 
sub-cylindrical,  and  bear  clusters  of  minor  pedicellariae  at  the  middle 
of  their  outer  sides.  There  are  four  rows  of  ventral  spines  (rarely 
five,  near  the  base  of  the  ray),  which  are  longer  than  the  ambula- 
crals,  slender,  with  acute  tips  pointing  outward.  At  the  bases  of  the 
ventral  spines  there  are  numerous  minor  pedicellariae,  clustered  at 
the  outer  side  in  the  inner  rows,  but  forming  wreaths  around  those 
of  the  outer  row.  The  marginal  row  of  dorsal  spines,  on  the  side  of 
the  ray,  consists  of  about  fifty  spines  as  slender  as  the  ventrals,  but 
capitate,  with  truncated  tips.  The  other  dorsal  spines,  above,  are  of 
two  kinds,  a  larger  and  a  smaller.  The  larger  ones  are  few  in  num- 
ber, shorter  but  much  thicker  than  the  ventrals,  capitate,  with  flat- 
tened heads,  and  are  arranged  in  a  pretty  regular  though  somewhat 
zigzag  median  row  of  about  twenty-five  spines,  crowded  near  the 
disc,  but  farther  apart  near  the  extremity  of  the  ray.  Between  this 
row  and  the  marginal  row  there  are  scattered  a  few  more  of  the 
larger  kind,  sometimes  in  clusters  or  short  rows  of  three  or  four.  On 
the  disc  they  form  a  more  or  less  distinct  pentagon,  within  which 


SHALLOW- WATER   STARFISHES  153 

there  is  another  circle,  and  a  spine  of  large  size  in  the  center.  The 
spines  of  the  smaller  kind,  minute,  slender,  and  truncated,  are  scat- 
tered between  the  large  ones.  Minor  pedicellariae  are  scattered  in 
considerable  numbers  between  the  spines,  and  form  wreaths  around 
the  bases  of  the  larger  ones.  The  major  pedicellariae  are  very  few  in 
number,  small  in  size,  and  of  rather  slender  form.  Papulae  numerous, 
but  not  forming  regular  groups.  Diameter,  five  inches.  (Stimpson.) 

"  A  pretty  and  well-characterized  species,  related  to  A.  epic  Mora. 

"  Habitat,  Puget  Sound.  North  West  Boundary  Commission.  Dr. 
C.  B.  Kennerly."  (Stimpson.) 

Two  nearly  typical  specimens  were  taken  by  the  Harriman  Expe- 
dition  at  Yakutat.  The  larger  of  these  is  a  dry  specimen  (a)  (pi. 
xxvi,  figs,  i,  2).  It  has  the  radii  18  mm.  and  120  mm.;  ratio, 
about  i :  6.6;  breadth  of  ray  at  base,  21  mm.  The  other  (&)  has  the 
radii  19  mm.  and  115  mm. ;  ratio,  about  i :  6. 

In  both  these  specimens  the  back  is  deeply  areolated  and  reticulated, 
the  spines  mostly  standing  in  single  rows  around  the  papular  areas, 
but  forming  clusters  at  the  principal  intersections  of  the  rows.  Both 
have  the  median  dorsal  band  of  spines  distinct  on  the  proximal  half 
of  the  ray,  but  quite  indistinct  distally,  where  the  areolations  become 
closer  and  the  spines  much  more  crowded  and  clustered.  The  spines 
in  both  are  short  and  capitate,  unequal  in  size,  the  larger  ones  sur- 
rounded by  a  cluster  of  smaller  ones,  but  in  one  specimen  (a)  the 
larger  spines  are  considerably  larger  than  in  the  other,  with  sub- 
truncate  rough  tips. 

In  one  specimen  (b)  the  center  of  the  disk  is  occupied  by  a  pretty 
regular  figure,  formed  by  five  polygonal  areas,  bordered  by  spines, 
and  surrounded  by  ten  other  polygonal  and  triangular  areas,  with 
others  at  the  radial  angles,  thus  forming  a  somewhat  stellate  figure. 
But  in  the  other  specimen  (a)  this  arrangement  is  obscured  by  the 
crowding  of  the  spines. 

The  superomarginal  spines  form  a  well  defined  band,  each  plate 
bearing  one  larger  capitate  spine,  like  the  dorsals,  and  two  to  five  or 
more  smaller  spines  of  the  same  kind.  There  is  a  well  defined  lateral 
channel  which,  in  one  of  these  specimens  (&)  bears  one  or  two  rows  of 
small  spines  distally,  besides  some  large  pedicellariae,  but  in  the  other 
(a)  these  interpolated  spines  are  almost  all  lacking. 

The  inferomarginal  row  of  spines  is  well  defined  and  nearly  regu- 
lar. In  one  case  (a)  each  plate  generally  bears  only  one  spine  proxi- 
mally,  but  on  the  distal  third  of  the  ray  there  are  mostly  two  equal 
spines,  close  together.  In  the  other  example  (&)  nearly  all  these 
plates  bear  two  stouter  and  blunter  spines,  to  the  base  of  the  rays. 


154  VERRILL 

On  the  proximal  part  of  the  rays  of  the  latter  there  are  three  rows 
of  strong  actinal  plates,  with  a  fourth  rudimentary  row  close  to  the 
base.  The  two  outer  rows  usually  bear  two  spines  to  a  plate;  the 
two  inner  ones  only  one  spine,  so  that  there  may  be  five  or  six  spines 
in  each  transverse  series.  But  in  the  other  specimen  (a)  the  larger 
proportion,  both  of  the  inferomarginal  and  actinal  plates,  bear  only 
one  spine,  so  that  the  total  number  is  much  less;  and  they  are  also 
less  stout,  a  little  tapered,  and  not  so  obtusely  rounded  at  the  tips, 
so  that  they  appear  much  more  openly  arranged  and  more  regu- 
lar. But  in  both,  these  ventral  spines  are  rather  large,  elongated, 
subequal,  sulcate  at  the  obtuse  tips,  often  bent  upward,  and  fre- 
quently compressed  when  crowded. 

Each  jaw  bears  a  pair  of  strong,  tapered,  blunt  teeth,  stouter  than 
the  adambulacrals,  and  an  external  strongly  divergent  pair,  about 
half  as  long;  on  the  external  end  of  the  jaw-plates  there  is  a  close 
pair  of  longer  and  more  tapered  spines.  The  next  four  or  five 
adambulacral  plates  (adorals)  are  closely  crowded  together,  and 
each  usually  bears  a  similar  long,  slender  adoral  spine.  In  many 
specimens  eight  to  ten  adoral  plates  are  monacanthid,  but  in  the 
largest  example  they  become  diplacanthid  at  about  the  fourth  to 
the  sixth,  varying  on  the  different  rays. 

The  more  distal  adambulacral  plates  are  pretty  regularly  dipla- 
canthid. The  two  spines  are  nearly  equal,  and  on  the  middle  of  the 
rays  are  about  equal  in  length  to  the  actinals ;  but  toward  the  mouth 
they  gradually  become  much  longer,  more  slender,  and  acute ;  most  of 
them  are  tapered  but  not  acute.  They  bear,  above  mid-height,  large 
clusters  of  forcipulate  minor  pedicellarise,  with  which  some  small 
forficulate  pedicellariae,  of  about  the  same  size,  are  often  inter- 
mingled. 

Thick  wreaths  of  clusters  of  minor  pedicellariae  also  occur  on  all 
the  ventral  and  upper  marginal  spines,  but  only  in  small  numbers  on 
the  dorsals.  They  also  occur  as  dermal  pedicellariae,  scattered  over 
the  dorsal  surface. 

Major  pedicellariae  are  few  on  some  specimens,  but  abundant  on 
others,  especially  on  the  lateral  channels  and  on  the  adoral  spines. 
The  largest  are  on  the  interradial  areas.  These  are  mostly  ovate- 
lanceolate,  or  regularly  lanceolate,  compressed,  and  have  very  acute 
tips.  Papulae  are  very  numerous. 

Color,  in  life,  is  often  bright  rosy  red  or  purplish. 

A  small  dried  specimen  (a)  from  Wrangel,  Alaska,  has  all  the 
essential  characters  of  the  larger  ones.  Its  radii  are  8  mm.  and 
.48  mm.  to  55  mm. ;  ratios,  1 : 6  or  1 :  7,  according  to  the  rays  measured. 


SHALLOW-WATER   STARFISHES  155 

The  rays  are  slender,  well  rounded  above,  and  regularly  tapered. 
The  dorsal  surface  is  covered  with  very  numerous  small  spines 
arranged  in  an  irregularly  reticulated  pattern,  much  as  in  the  adult, 
but  on  a  much  smaller  scale.  The  spines  are  capitate  and  unequal. 

The  upper  and  lower  marginal  spines  and  two  rows  of  actinals 
stand  in  simple  regular  rows,  one  to  a  plate.  The  peractinal  row 
extends  to  the  tips  of  the  arms,  the  spines  becoming  very  small  dis- 
tally ;  the  first  row  of  subactinals  only  extends  to  about  the  middle  of 
the  rays;  a  few  spines  occur  on  a  rudimentary  synactinal  series  at 
the  base  of  the  arms. 

The  adambulacral  plates  usually  bear  alternately  one  and  two 
slender  spines.  The  minor  pedicellarise  occur  in  clusters  on  nearly 
all  the  spines ;  major  pedicellariae  are  very  few  and  small. 

Specimens  considerably  smaller  than  this  are  similar  in  character 
and  could  hardly  be  mistaken  for  any  other  species. 

Two  small  specimens  (d,  e)  were  taken  by  Dr.  Coe  at  Sitka  and 
preserved  in  alcohol.  The  larger  (d)  (pi.  xxii,  figs.  I,  2)  has  the 
radii  12  mm.  and  55  mm.  The  dorsal  spines  show  above  the  general 
surface  only  slightly,  their  rounded  tips  alone  being  visible,  but  their 
clustered  arrangement  is  apparent,  and  gives  the  surface  an  irregular, 
roughly  uneven  appearance.  The  wreaths  of  minor  pedicellariae  are 
dense  and  attached  near  the  tips  of  the  spines,  and  with  the  abun- 
dant papulae  they  closely  cover  all  the  spaces  between  the  spines. 

The  marginal  and  ventral  spines  are  more  conspicuous,  their  large 
clusters  of  pedicellariae  being  attached  lower  down.  These  spines 
form  four  very  regular  longitudinal  rows,  the  upper  marginals  being 
smaller  than  the  others.  The  adambulacral  spines  are  almost  con- 
cealed by  their  large  clusters  of  pedicellariae. 

The  smaller  example  (e)  has  the  radii  5  mm.  and  40  mm.  The 
rays  are  round  and  slender,  rather  rigid,  and  taper  gradually.  The 
surface  and  spines  agree  with  the  preceding,  except  that  the  dorsal 
spines  are  even  less  apparent  and  the  surface  less  uneven,  though 
the  clustered  arrangement  of  the  spines  is  still  evident. 

The  color  of  these  alcoholic  specimens  is  light  rusty-brown.  In 
life  it  is  usually  light  red. 

VARIATIONS. 

A  larger  specimen  (f)  from  Puget  Sound  (Kincaid)  has  the  radii 
22  mm.  and  162  mm. ;  ratio,  about  1 : 7.3.  The  five  rays  are  long, 
well  rounded  and  gradually  tapered.  It  agrees  pretty  well  with 
those  described  above,  but  the  reticulations  made  by  the  dorsal  spines 


156 


VERRILL 


are  less  complete  and  less  regular,  both  on  the  rays  and  on  the  disk. 
Radial  bands  of  spines  are  not  distinct.  The  spines  are  also  smaller 
and  less  decidedly  capitate. 

The  inferomarginal  and  actinal  spines  mostly  stand  singly,  so  that 
there  are  usually  only  four  or  five  rows  near  the  base  of  the  rays, 
but  on  some  of  the  rays  more  or  less  of  the  plates  bear  two  spines. 

The  adambulacral  spines  are  long  and  slender,  about  equal  in 
length  to  the  actinals  on  the  greater  part  of  the  length  of  the  rays  ; 
but  toward  the  mouth  they  increase  in  length  more  rapidly  than  the 
actinals,  becoming  decidedly  long  and  slender  on  the  adoral  plates. 

Major  pedicellariae  are  abundant  in  the  lateral  furrows,  interradial 
areas,  between  the  actinal  spines,  in  clusters  on  the  adambulacral 
spines,  and  also  on  the  inner  edges  of  the  ambulacral  plates.  They 
are  of  moderate  size,  compressed,  mostly  acute-ovate  or  ovate-lan- 
ceolate in  form.  Those  on  the  sides  and  bases  of  the  rays  are  the 
larger.  The  clusters  on  the  adambulacral  spines  are  in  many  cases 
composed  of  small-sized  forficulate  major  pedicellariae,  but  on  other 
spines  they  are  mixed  with  forcipulate  minor  pedicellariae  in  varying 
proportions. 

Minor  pedicellariae  occur  in  small  numbers  on  nearly  all  the 
spines.  They  are  also  abundant  between  the  dorsal  spines,  on  the 
papular  areas,  and  on  the  surfaces  of  the  lateral  channels.  They 
generally  stand  singly  when  dermal.  The  papulae  are  small  and  very 
numerous,  in  large  clusters. 

In  the  Museum  of  Comparative  Zoology  I  have  studied  a  good 
series,  including  some  that  are  quite  small. 

One  series  from  Mendocino,  California,  includes  specimens  only 
35  mm.  to  40  mm.  in  diameter  (Nrx  llffi).  Lot  No.  1192,  also  from 
California,  includes  three  young.  No.  1195  is  from  Alaska.  Nos. 
1^  1187  are  typical  specimens  from  the  Gulf  of  Georgia 
(coll.  A.  Agassiz).  Of  these,  No.  1191,  was  labelled  as  Asterias 
brachiata  of  Perrier.  It  is  probably  a  cotype  of  his  species,  but  is 
not  the  one  of  which  he  gave  measurements.  He  gave  no  locality 
for  his  species.  There  can  be  no  doubt  that  the  type  was  from  this 
lot  and  that  it  is  a  synonym  of  E.  troschelii  (Stimpson).  No.  1190 
is  a  short-rayed,  close-spined  variety  from  Mendocino,  California, 

No.  1906,  from  Friday  Harbor,  Puget  Sound  (Professor  Kincaid), 
is  a  short-rayed  specimen  with  coarse  spinulation  (radii,  22  mm.  and 
140  mm.). 

The  dorsal  spines  are  decidedly  longer  than  usual,  capitate,  con- 
spicuously reticulated  and  areolated,  with  no  distinct  median  radial 


'' 
1  1  ^  3 


SHALLOW-WATER   STARFISHES 


rows.  Thus  its  dorsal  spinulation  is  similar  in  appearance  to  that  of 
some  examples  of  Pisaster  ochraceus.  But  the  stout  marginal  spines 
and  three  regular  rows  of  actinals  are  arranged  nearly  as  in  the 
typical  troschelii,  and  its  adambulacrals  are  irregularly  diplacanthid. 
No  large,  unguiculate  dorsal  pedicellariae  were  found.  Although  it 
looks  like  a  hybrid  with  ochraceus,  it  is  near  the  type  of  var.  alve- 
olata. 

The  dermal  major  pedicellariae  are  small,  lanceolate,  pointed. 
Minute  minor  pedicellariae  are  sparsely  scattered  on  the  integument 
of  the  back,  and  form  thin  wreaths  around  the  spines,  but  on  the 
adambulacral  and  actinal  spines  they  are  abundant,  forming  distal 
clusters,  in  which  there  are  also  a  few  small,  scattered,  pointed 
major  pedicellariae. 

The  papular  areas  on  the  dorsal  side  are  large  and  the  papulae 
very  numerous,  in  large  groups;  between  the  actinal  and  marginal 
rows  they  are  large  but  in  small  clusters. 

Some  other  specimens  of  this  species  have  the  rays  much  shorter 
than  usual.  This  is  the  case  with  Nos.  1421  and  1432  (Mus.  Comp. 
Zool.)  from  the  Gulf  of  Georgia.  In  these  the  dorsal  spines  are 
numerous  and  crowded,  areolated  or  reticulated.  They  somewhat 
resemble  var.  alveolata,  but  the  shortness  of  the  rays  may  perhaps 
be  due  to  mutilation  and  imperfect  restoration,  for  the  rays  are  not 
precisely  equal. 

One  of  the  larger  specimens  (about  twenty  inches  in  diameter) 
studied  by  me  is  from  Esquimault,  B.  C.  (Prov.  Mus.  B.  C.),  sent  by 
Dr.  Newcombe. 

The  radii  are  43  mm.  and  250  mm. ;  ratio,  1 :  5.8 ;  breadth  of  rays 
near  the  middle,  68  mm.  to  76  mm. 

The  disk  and  rays  were  evidently  distended  with  air  before  drying, 
as  shown  by  the  wide  inflated  form,  and  the  widely  stretched  dorsal 
papular  areas;  but  near  the  tips  of  the  rays  the  normal  rounded 
form  is  retained,  with  the  close  reticulate  arrangement  of  the  spines. 

The  dorsal  spines  are  mostly  small  and  acute,  standing  on  the 
openly  reticulated  ossicles  in  short  rows,  the  transverse  rows  being 
most  conspicuous,  especially  on  sides  of  the  rays.  But  there  is 
a  slightly  distinct  medial  row  of  somewhat  larger  and  partly  capitate 
spines  on  the  proximal  half  of  the  rays.  The  disk  is  coarsely  reticu- 
lated with  slightly  larger  clavate  and  capitate  spines.  The  madre- 
porite  is  large,  complicated,  but  not  surrounded  by  any  special  spines 
or  grooves. 

Notwithstanding  its  great  size,  the  number  of  rows  of  marginal 
and  actinal  spines  is  not  greater  than  in  some  specimens  of  var.  sub- 


150  VERRILL 

nodosa,  not  more  than  one-fourth  its  diameter.  The  upper  and 
lower  marginals  are  wide  apart ;  the  upper  row  is  mostly  single ;  the 
lower,  mostly  double ;  the  peractinal  and  the  first  subactinal  row  are 
double;  the  imperfect  synactinal  row  is  single  so  that  there  are 
about  eight  pretty  regular  and  even  ventral  rows  proximally.  All 
these  spines  and  the  adambulacrals  are  shaped  as  in  the  specimens 
already  described. 

Rather  large,  acute-ovate,  dermal  major  pedicellariae  are  numerous 
between  the  actinal  and  marginal  rows,  and  large  clusters  of  smaller 
ones  are  abundant  on  the  adambulacral  spines.  Papulae  are  exceed- 
ingly numerous  in  large  groups  on  the  dorsal  and  lateral  areas. 

The  Asterias  epichlora  of  M.  de  Loriol  (non  Brandt,  op.  cit, 
1897)  is  doubtless  identical  with  this  species,  and  is  not  the  true 
epichlora  of  Brandt,  which  was  evidently  the  common  small  greenish 
Sitka  species,  and  probably  identical  with  A.  saanichensis  De  Loriol 
(op.  cit.,  p.  23,  pi.  n,  figs.  3-3^  4,  5).  This  has  been  discussed  under 
A.  epichlora. 

The  differences  that  he  found,  in  comparison  with  Stimpson's 
description  of  troschelii,  are  variable  characters  in  this  species,  as  my 
descriptions  above  will  show.  Some  of  the  specimens  studied  by  me 
agree  well  with  the  one  figured  by  M.  de  Loriol  and  came  from  the 
same  district. 

The  geographical  range  of  this  species  is  extensive.  I  have 
studied  specimens  from  Mendocino,  California,  and  Yakutat,  Alaska. 
It  was  taken  by  the  Harriman  Expedition  on  the  Alaskan  coast,  at 
Sitka,  Wrangel,  Orca,  and  Yakutat.  Mr.  J.  F.  Whiteaves,  of  the 
Canadian  Geological  Survey,  sent  me  specimens  from  the  Straits 
of  Georgia,  Malaspina  Inlet,  Discovery  Passage,  and  Queen  Char- 
lotte Islands.  Dr.  C.  F.  Newcombe  sent  many  specimens  from  Vic- 
toria and  Esquimault  Harbor,  some  of  large  size.  I  have  also 
examined  specimens  from  Puget  Sound  (Kincaid)  and  several  other 
localities.  Departure  Bay,  British  Columbia  (H.  J.  Young,  Canadian 
Geological  Survey,  1908),  many  large  and  small;  Gulf  of  Georgia 
(A.  Agassiz,  Mus.  Comp.  Zool.),  a  good  series. 

EVASTERIAS  TROSCHELII  Var.  RUDIS  Verrill,  nov. 

The  three  large  specimens  (h,  i,  /)  about  equal  in  size  and  form, 
from  British  Columbia,  appear  to  belong  to  this  species,  though  they 
have  larger  and  swollen  rays,  and  much  coarser  dorsal  spines.  One 
of  these  has  the  radii  36  mm.  and  252  mm. ;  ratio,  1 : 7.  The 
specimens  are  flattened  in  drying,  the  dorsal  skeleton  being  weak, 
and  this  makes  the  rays  appear  broader. 

«<•"' 
I     t 

P/VA.A'  U|M       <**>** 


SHALLOW- WATER   STARFISHES  159 

The  dorsal  surface  is  very  openly  reticulated,  with  rather  small 
and  narrow  ossicles,  forming  large,  irregular,  angular  papular  areas. 

The  dorsal  spines  are  rough,  unequal,  and  numerous,  though  less     \*A\M/ 
numerous  than  in  the  type.    They  are  irregularly  and  coarsely  reticu- 
late in  arrangement ;  and  toward  the  base  of  the  rays,  especially  on 
the  sides,  the  smaller  ones  form  short  transverse  rows  or  combs  on 
the  borders  of  the  larger,  transversely  elongate,  papular  areas. 

The  larger  spines,  which  are  relatively  few  in  number,  are  short, 
strongly  clavate  or  subtruncate,  and  striate  at  the  tips,  mostly  scarcely 
higher  than  broad.  These  larger  spines  often  form  an  irregular 
median  radial  row  proximally;  others  are  irregularly  placed  on  the 
nodes  of  the  larger  reticulations.  Toward  the  ends  of  the  rays,  they 
become  much  more  numerous  and  sometimes  stand  in  small  groups. 

The  smaller  spines  are  very  much  smaller,  mostly  tapered  and  sub- 
acute,  but  some  are  obtuse  and  many  are  spinulose  at  the 
tip.  The  whole  dermal  surface,  including  the  papular  areas,  is  cov- 
ered with  numerous  small  minor  pedicellariae.  They  also  form  small 
clusters  on  the  bases  of  the  spines.  The  superomarginal  spines  are 
rather  longer  than  the  dorsals  and  tapered  or  only  slightly  clavate, 
often  obliquely  truncate ;  they  mostly  stand  singly  on  the  plates  and 
form  a  regular  row,  but  on  one  specimen  they  stand  two  or  even  i 
three  on  a  plate,  the  secondary  ones  being  smaller. 

The  inferomarginals  are  very  similar  in  size  and  form,  but  rather 
more  tapered;  they  usually  stand  two  on  a  plate,  but  sometimes 
singly  proximally. 

There  are  usually  three  rows  of  actinal  ossicles  proximally,  some- 
times four,  rarely  but  two.  Many  of  them  bear  two  or  even  three 
spines,  others  but  one,  so  that  the  number  of  rows  of  actinal  spines 
is  variable;  but  they  are  always  numerous,  crowded,  nearly  equal, 
and  very  similar  to  the  inferomarginals,  though  usually  rather 
longer  and  more  tapered,  especially  toward  the  mouth. 

The  adambulacral  spines  are  terete,  much  more  slender,  but  about 
as  long  as  the  actinals,  they  mostly  stand  alternately,  one  and  two  to 
a  plate.  Those  within  the  disk,  and  especially  the  adorals  and  epi- 
orals,  become  decidedly  longer  (6  mm.  to  8  mm.)  and  more  slender. 
The  apical  perorals  are  much  shorter  and  stouter. 

The  lateral  channels,  between  the  upper  and  lower  marginal 
spines,  and  also  between  the  latter  and  the  peractinals,  are  well 
defined  and  bear  numerous  pedicellariae  of  both  kinds. 

The  major  pedicellariae  are  acute-ovate  or  triangular-ovate,  acute 
at  the  tip,  and  not  very  large.  They  also  occur  on  the  back  and 


l6o  VERRILL 

among  the  ventral  and  adambulacral  spines.  The  latter  bear  dense 
clusters  of  minor  pedicellariae  on  the  outer  surface,  and  near  their 
tips. 

The  madreporic  plate  is  large,  with  numerous  fine,  forked,  radial 
gyri.  It  is  not  surrounded  by  a  special  circle  of  spines. 

Another  large  specimen  (fc)  from  Victoria,  British  Columbia, 
differs  considerably  in  the  spinulation,  especially  on  the  ventral  side. 

The  radii  are  36  mm.  and  232  mm. ;  ratio,  i :  6.45. 

The  rays  are  stout  and  broad,  but  unnaturally  flattened  in  drying, 
so  that  they  taper  rather  abruptly  distally. 

The  smaller  dorsal  spines  are  numerous,  mostly  in  reticulated 
lines  and  in  short  transverse  rows  or  combs,  the  transverse  lines 
being  most  conspicuous  proximally  and  toward  the  sides  of  the  rays, 
dorsally;  not  so  evident  on  the  disk.  These  smaller  spines  mostly 
stand  in  a  single  row  on  the  narrow  carina  of  the  ossicles  surround- 
ing the  large  papular  areas.  Part  are  slender  and  acute,  about  I  mm. 
to  1.5  mm.  long  and  0.25  mm.  thick.  More  than  half  of  them  are 
obtuse  or  clavate,  of  about  the  same  length,  but  from  0.40  mm.  to 
0.60  mm.  thick. 

Larger  capitate  and  sulcated  spines  are  scattered  among  the 
smaller  reticulated  spines.  They  stand  on  the  larger  ossicles,  and 
especially  along  the  median  radial  lines,  forming,  with  many  smaller 
ones,  rather  irregular  radial  ranges  of  spines.  On  the  distal  part  of 
the  rays  the  spines  become  more  crowded  and  the  larger  ones  more 
numerous,  and  are  then  often  grouped  more  or  less  in  clusters. 

The  madreporic  plate  is  large,  with  fine  radial  gyri,  and  is  sur- 
rounded by  an  irregular  row  of  small  spines.  Minor  pedicellariae 
are  thickly  scattered  over  the  papular  areas  and  surround  all  the 
spines,  in  small  wreaths. 

The  superomarginal  spines  form  a  pretty  regular  row,  in  which 
the  larger,  blunt  spine  on  each  plate  is  accompanied  by  one  to  three 
smaller,  more  acute  ones.  The  ventral  spines  usually  consist  of  three, 
often  double,  rows  and  a  single  synactinal  row,  all  crowded  pretty 
closely  together,  and  nearly  equal  in  length,  but  differing  in  form. 

Most  of  the  inferomarginal  ossicles  bear  two  equal  spines  and 
sometimes  three,  so  that  there  may  be  from  four  to  seven  spines  in 
each  transverse  range.  These  spines  are  mostly  rather  short,  stout, 
partly  blunt,  partly  tapered,  often  bent,  some  of  those  toward  the 
mouth  becoming  longer,  subacute,  and  often  excurved.  They  are 
mostly  about  I  mm.  thick  and  2  mm.  to  3  mm.  high. 


SHALLOW-WATER   STARFISHES  l6l 

The  ventral  ossicles  are  stout  and  tesselated;  in  size,  shape,  and 
number  very  nearly  as  in  the  typical  form.  The  adambulacral  spines 
are  longer  and  more  slender  than  in  the  type,  especially  toward  the 
mouth,  where  they  increase  more  in  length,  some  of  the  adorals 
becoming  6  mm.  to  7  mm.  long.  They  are  mostly  alternately  one 
and  two  to  a  plate.  The  apical  peroral  spines  are  stout  and  blunt, 
rather  long.  The  dermal  major  pedicellariae  are  nearly  as  in  the 
type,  but  some  are  larger  and  stouter.  There  are  few  on  the  dorsal 
surface. 

This  large  and  peculiar  variety  might  well  be  thought  a  distinct 
species,  had  there  not  been  specimens  more  or  less  intermediate  in 
the  collections  from  Victoria  and  the  Queen  Charlotte  Islands.  It  may 
be  that  it  is  the  condition  commonly  assumed  when  grown  to  unusual 
size. 

In  form,  size  and  general  appearance  this  variety  resembles 
E.  accmthostoma,  but  the  dorsal  spines  of  the  latter  are  much 
smaller  and  more  acute,  and  form  conspicuous  transverse  combs. 

Victoria  and  Esquimault  Harbor  (Prov.  Mus.  B.  C,  collected  by 
Dr.  C.  F.  Newcombe)  ;  Queen  Charlotte  Islands  (Canadian  Geolog- 
ical Survey). 

EVASTERIAS  TROSCHELII  Var.  DENSA  Verrill,  nov. 

One  young  specimen  (g)  from  Victoria,  British  Columbia,  is 
peculiar  in  having  the  dorsal  spines  all  nearly  uniform  in  size  and 
form,  all  being  capitate  and  arranged  in  a  rather  close  reticulate 
pattern,  the  spines  standing  in  single  rows  and  nearly  in  contact  on 
the  reticulating  ossicles.  Proximally  they  often  form  short  trans- 
verse rows  on  the  sides  of  the  rays,  but  distally  they  are  rather  uni- 
formly crowded  and  not  acervate,  nor  is  there  any  distinct  median 
radial  row,  nor  any  larger  primary  spines  elsewhere. 

The  reticulations  are  much  smaller,  more  transverse,  and  closer 
than  in  the  typical  form;  this,  and  the  absence  of  pedicellariae  in 
clusters  around  the  large  spines,  give  it  a  very  different  appearance, 
approaching  that  of  the  large  specimen  of  E.  acanthostoma,  in  this 
respect. 

There  are  five  regular  rows  of  convex,  imbricated,  ventral  ossicles 
proximally,  many  of  which  bear  two  spines,  which  are  crowded  and 
much  like  those  of  the  larger  form,  but  rather  more  slender  and  less 
obtuse. 

Radii,  13  mm.  and  78  mm. ;  ratio,  i :  6. 

Near  Victoria,  British  Columbia  (Prov.  Mus.  B.  C.). 

12 


l62  VERRILL 

EVASTERIAS  TROSCHELII  Var.  ALVEOLATA  Verrill,  nov. 

Plate  LXII,  figure  i. 

A  five-rayed,  conspicuously  reticulated  variety,  with  plump, 
rounded,  tapered,  subacute  rays  of  medium  length.  Radii  of  the 
type,  22  mm.  and  no  mm. ;  ratio,  1:5.  Madreporite  large,  round, 
convex,  yellow,  surrounded  by  a  circle  of  fine  capitate  spines. 

The  dorsal  ossicles  are  narrow,  but  high,  with  subacute  summits, 
on  which  are  rows  of  small,  slender,  capitate  spines.  They  surround 
rather  deeply  depressed,  angular,  papular  areas,  variable  in  size  and 
form,  but  often  hexagonal  or  pentagonal;  larger  and  more  rectan- 
gular ones  are  on  the  sides  of  the  rays,  between  the  marginals. 

The  dorsal  spines  are  somewhat  unequal,  the  larger  ones  standing 
at  the  intersections  of  the  ossicles  and  on  the  median  line,  but  forming 
only  a  very  inconspicuous  median  band ;  they  are  not  at  all  acervate. 

The  marginal  and  actinal  spines  form  five  nearly  simple,  regu- 
larly spaced  rows  proximally,  all  of  which  are  similar  in  size  and 
form,  except  the  upper  marginals,  which  are  shorter,  thicker,  and 
more  capitate.  These  mostly  stand  one  to  a  plate,  but  sometimes 
two ;  or  there  may  be  one  or  two  small  accessory  spines  on  the  plate, 
or  on  the  connecting  transverse  ossicles.  The  spines  of  the  other 
rows  are  a  little  longer,  short-clavate  or  obtuse,  and  mostly  one  to  a 
plate. 

The  peractinal  and  subactinal  rows  have  one  or  two  to  a  plate,  or 
alternately  one  and  two  in  places.  The  synactinal  row  extends  to 
about  the  distal  fourth  of  the  ray. 

The  adambulacral  spines  stand  mostly  two  to  a  plate;  they  are 
rather  stout,  cylindrical,  or  slightly  tapered,  acute. 

Pedicellarise  are  few.  Very  small,  ovate  dermal  major  pedicel- 
lariae  are  scattered  on  the  dorsal  side ;  and  larger  ones,  more  acute 
in  form,  are  found  on  the  interradial  and  marginal  areas. 

The  color,  as  dried,  is  dull  purple. 

Departure  Bay,  British  Columbia  (Canadian  Geological  Survey, 
1908). 

This  looks  like  a  distinct  species,  but  is  probably  a  short-rayed 
variety  of  E.  troschelii.  Its  rays  are  not  only  shorter  and  thicker, 
but  the  areolations  are  deeper  and  more  sharply  defined,  while  the 
marginal  and  actinal  spines  are  larger  and  fewer,  and  the  rows  are 
fewer  and  more  widely  separated.  It  may,  however,  eventually  prove 
to  be  a  distinct  species,  when  a  more  extensive  series  can  be  studied. 


SHALLOW-  WATER   STARFISHES  163 

EVASTERIAS  TROSCHELII  Var.  SUBNODOSA  Verrill,  nov. 

Several  specimens,  mostly  of  rather  small  size,  differ  from  the 
ordinary  variety  so  much  in  appearance  that  they  seem  to  deserve  a 
varietal  name,  though  intermediate  specimens  occur.  In  its  extreme 
form  small  specimens  of  this  variety  closely  resemble  some  five-rayed 
varieties  of  L.  epichlora,  especially  on  the  upper  side.  Others, 
70  mm.  to  80  mm.  in  diameter,  so  closely  resemble  dorsally  the  young  *  - 

of  Pisaster  ochraceus  as  to  be  readily  mistaken  for  that  species  ;  but  - 
they  can  easily  be  distinguished  by  the  subdiplacanthid  adambulacral 


spines,  slender  actinal  spines,  and  the  small  pedicellariae.  j> 

The  special  features  of  this  variety  are  the  conspicuously  clustered 


or  somewhat  acervate  arrangement  of  the  dorsal  spines,  some  of 
which  are  larger  and  thicker  than  usual,  with  strongly  capitate  and 
more  or  less  flat-topped  tips,  while  there  are  minute  rows  of  much 
smaller  clavate  and  capitate  spines  scattered  between  them  and  partly 
on  the  skeletal  network  ;  the  very  smallest  are  often  acute. 

All  of  the  spines,  above  and  below,  bear  large  clusters  of  minor    ^^  &f 
pedicellariae,  and  small  dermal  clusters  are  abundantly  scattered  on 
the  back,  so  that,  in  alcohol,  the  dermis  and  most  of  the  smaller 
spines  are  quite  concealed  by  them,  and  the  papulae.  /l/Ori-i**- 

The  two  rows  of  marginal  and  two  to  four  rows  of  actinal  spines 
are  pretty  uniform  in  diameter  and  regular,  but  usually  crowded,  in  . 
arrangement;  they  are  nearly  as  stout  as  the  larger  dorsals  and 
somewhat  longer,  the  inferomarginals  and  actinals  being  longer  than 
the  superomarginals,  and  a  little  curved  upward.  They  are  mostly  -CA^^MMP  < 
a  little  clavate  and  obtuse.  In  the  larger  examples  the  inferomar- 
ginal  and  actinal  rows  become  double,  so  that  there  may  be  six  or 
seven  rows.  The  adambulacral  spines  are  crowded,  terete,  obtuse, 
becoming,  as  usual  in  all  varieties  of  this  species,  distinctly  longer 
and  more  slender  toward  the  mouth  ;  they  are  loaded  with  clusters 
of  small  pedicellariae  and  are  irregularly  diplacanthid. 

One  typical  specimen  from  Wrangel,  Alaska,  has  the  radii  10  mm. 
and  60  mm.  ;  ratio,  1  :  6.  Another  from  Victoria,  Vancouver  Island 
(Prov.  Mus.  B.  C),  has  the  radii  22  mm.  and  137  mm.;  ratio, 
i  :  6.23. 

EVASTERIAS  TROSCHELII  Var.  PARVISPINA  Verrill,  nov. 

Plate  cvi,  figures  i,  2. 

Rays  five,  in  the  type  ;  firm,  rather  short,  round  and  plump,  taper- 
ing rather  rapidly  distally.  Radii,  6  mm.  and  28  mm.  The  whole 
upper  surface  in  alcoholic  specimens  is  nearly  uniformly  concealed 


164  VERRILL 

by  the  pedicellariae  and  papulae,  so  that  the  small,  uniform  spines  are 
scarcely  visible,  and  there  is  no  apparent  grouping  into  clusters,  nor 
any  evident  rows.  The  clusters  of  minor  pedicellariae  are  dense, 
though  not  large,  but  they  are  mostly  in  contact  with  their  neighbors, 
owing  to  the  numerous  spines.  They  seem  to  be  attached  close  to 
the  tips  of  the  spines. 

The  lateral  and  ventral  spines  form  about  four  distinct,  regular, 
but  crowded,  evenly  spaced  rows,  and  are  rather  more  distinctly 
defined  than  the  dorsals;  but  like  the  adambulacrals,  they  are  well 
covered  by  large  clusters  of  minor  pedicellariae. 

Color  rusty-brown,  as  preserved. 

When  dry,  the  dorsal  surface  is  covered  with  very  numerous,  but 
not  closely  crowded,  small,  unequal  spines,  arranged  in  small  groups 
or  singly,  so  as  to  form  irregular  circles  around  the  papular  areas, 
thus  giving  the  surface  an  imperfectly  and  irregularly  areolated 
appearance.  The  median  radial  spines  are  a  little  larger,  forming  an 
irregular  crooked  row.  The  dorsal  spines  are  all  short,  their  height 
scarcely  twice  their  thickness,  in  the  larger  ones,  but  the  smaller 
ones  are  much  more  slender.  The  larger  are  capitate,  with  rough 
rounded  tips;  others  are  clavate.  The  papular  areas  are  numerous, 
round,  rather  small,  but  unequal  in  size.  The  papulae  are  unequal, 
and  mostly  stand  singly  or  in  small  groups. 

The  lateral  and  ventral  spines  are  all  a  little  capitate,  shaped  like 
the  larger  dorsals,  but  a  little  longer  and  larger.  They  form  five 
pretty  regularly  spaced,  simple  rows  (sometimes  six)  on  the  proxi- 
mal three-fourths  of  a  ray,  separated  by  four  rows  of  papulae.  Four 
of  these  rows  of  spines  extend  nearly  or  quite  to  the  tips  of  the  rays, 
where  they  become  much  crowded.  The  upper  marginal  series  is 
usually  double  proximally,  with  two  spines  on  a  plate,  but  single 
distally.  A  narrow  channel  with  a  regular  row  of  round  papular 
areas  separates  them  from  the  lower  marginals.  These  form  a 
regular  row  to  the  tips;  it  is  double  on  the  basal  part,  two  spines 
standing  close  together  on  most  of  the  plates,  but  distally  it  appears 
double  by  the  crowding  of  the  row  next  below.  The  latter  is  sepa- 
rated on  the  basal  third  of  the  ray  by  a  row  of  small  papulae;  but 
these  are  absent  distally,  so  that  the  spines  seem  to  belong  to  the 
lower  marginal  series,  as  both  rows  extend  to  the  tips  of  the  rays, 
but  they  are  probably  true  peractinals.  These  are  followed  by  a 
row  of  papulae,  and  then  by  a  row  of  subactinals  that  extends  to 
about  the  middle  of  the  ray.  All  these  ventral  spines  are  much  alike 
in  size  and  form,  and  they  are  pretty  regularly  and  evenly  spaced. 


SHALLOW-WATER   STARFISHES  165 

The  adambulacral  spines  are  crowded  in  two  or  three  irregular 
rows,  the  plates  bearing  either  one  or  two  in  irregular  alternation. 
They  are  about  as  long  as  the  peractinal  spines  and  resemble  them 
in  form,  but  are  more  slender.  They  are  mostly  clavate  with  rounded 
rough  tips.  The  outer  ones  are  the  larger.  They  bear  small 
clusters  of  pedicellariae.  Minor  pedicellariae  also  occur  in  small, 
dense  clusters  on  all  the  ventral  and  dorsal  spines,  and  singly  on  the 
dorsal  papular  areas ;  they  are  very  small. 

Major  pedicellariae  are  very  few.  The  largest  are  relatively  large, 
nearly  equal  to  the  adjacent  small  spines  in  thickness,  stout,  triangu- 
lar-ovate, obtuse;  these  stand  erect  in  the  actinal  interradial  areas. 
A  few  very  much  smaller  ones,  similar  in  shape,  occur  on  the  adam- 
bulacral spines. 

The  dermal  ossicles  are  numerous  and  unusually  well  developed, 
so  that  the  dried  specimen  is  firm  and  rigid. 

Sitka  (Dr.  W.  R.  Coe,  Harriman  Expedition). 

These  specimens  superficially  somewhat  resemble  C.  cribraria,  but 
are  very  distinct  in  the  form  and  size  of  the  pedicellariae,  in  the  much 
stouter  dorsal  ossicles,  in  the  arrangement  of  the  dorsal  spines,  and 
in  the  form  of  the  marginal,  ventral,  and  adambulacral  spines. 

This  is  quite  unlike  all  the  varieties  of  L.  epichlora  in  appearance. 
It  was  at  first  thought  to  be  a  new  species.  It  may  be  distinguished 
from  epichlora  by  the  small  size  of  the  dorsal  spines  and  their 
areolate  arrangement ;  they  are  not  crowded  as  in  subsp.  miliaris.  If 
they  were  all  enlarged  the  appearance  would  be  much  as  in  subsp. 
alaskensis.  The  numerous  regular,  even  rows  of  slender  marginal 
and  actinal  spines  are  also  characteristic.  The  dorsal  ossicles  are 
numerous  and  form  a  strong  reticulation.  No  very  large,  serrate 
major  pedicellariae  occur  on  the  types. 

This  is,  no  doubt,  one  of  the  regular  young  forms  of  E.  troschelii, 
but  my  specimens  are  not  sufficiently  numerous  to  connect  it  with 
either  of  the  special  varieties  of  that  species.  It  is  apparently  most 
like  the  typical  form. 

EVASTERIAS  ACANTHOSTOMA  Verrill,  sp.  nov. 
Plate  xx,  figures  I,  a;  plate  xxiv,  figure  3  (type). 

A  large  species  with  a  rather  contracted  disk  and  five  long,  tapered, 
flexible  arms.  Lesser  radii  of  the  type,  30  mm. ;  greater  radii,  275 
mm.  Ratio,  as  1:9. 

It  is  remarkable  for  the  length  of  the  oral  and  adoral  spines  and 
the  adjacent  actinal  spines ;  for  the  numerous  crowded  ventral  spines, 


l66  VERRILL 

which  form  six  to  eight  rows;  for  the  numerous  very  small  dorsal 
spines,  mostly  arranged  in  small  transverse  rows  or  combs;  and  for 
the  deep  ambulacral  grooves  and  long  ambulacral  feet. 

The  adambulacral  spines  are  crowded  in  three  irregular  rows, 
most  of  them  standing  two  on  a  plate,  but  frequently  alternating  with 
one  on  a  plate  (subdiplacanthid).  Those  at  and  near  the  mouth 
are  very  long  and  rather  slender,  terete,  slightly  fusiform,  pointed. 
The  longest  are  up  to  8  mm.  or  9  mm.  long.  They  decrease  in 
length  distally  and  become  more  obtuse;  about  opposite  the  border 
of  the  disk  they  are  only  about  half  as  long  (4  mm.  to  5  mm.),  or 
about  equal  in  length  to  the  adjacent  actinal  spines,  but  are  smaller 
and  more  tapered.  They  all  bear  dense  clusters  of  minor  pedicel- 
lariae  near  the  ends. 

The  interactinal  ossicles  form  about  five  regular  close  rows,  nearly 
equal  in  size  and  form.  The  synactinal  ones  are  only  a  little  smaller, 
and  like  the  others  may  bear  two  or  three  spines.  Many  of  the 
actinal  ossicles  bear  two  spines  that  stand  obliquely  on  the  ossicle; 
hence  there  are  six  to  eight  rows  of  crowded,  nearly  equal  spines, 
which  are  strong,  not  very  long,  mostly  somewhat  fusiform,  and 
tapered  distally,  obtuse  or  subtruncate,  often  slightly  compressed, 
substriate  or  nearly  smooth,  mostly  slightly  curved  outward.  They 
bear  small,  dense  clusters  of  minor  pedicellariae  on  the  outer  side,  or 
sometimes  small  wreaths.  Between  all  the  actinal  plates  are  regular 
papular  areas,  with  clusters  of  small  papulae.  Toward  the  base  of 
the  arms,  and  especially  on  the  under  side  of  the  disk,  these  spines 
become  longer  and  more  acute,  some  of  the  most  proximal  becoming 
twice  the  length  of  those  farther  out.  The  rows  of  ossicles  diverge 
somewhat  at  the  base  of  the  rays,  and  the  outer  ones  curve  upward. 

Separated  from  the  inferomarginals  or  outer  ventral  row  by  a 
very  distinct  papular  channel,  there  is  a  row  of  strong  superomar- 
ginal  ossicles,  each  of  which  usually  bears  two  or  three  spines,  simi- 
lar to  the  actinal  spines  in  length,  but  smaller  and  rougher,  with 
sulcated  tips.  This  row  of  ossicles  and  spines  curves  upward  to  the 
dorsal  surface  at  the  base  of  the  rays,  and  there  meets  the  corre- 
sponding row  of  the  next  ray. 

The  dorsal  spines,  which  are  very  small,  nearly  equal,  either 
tapered  or  clavate,  with  rough  sulcated  tips,  are  very  numerous, 
without  any  regular  arrangement.  They  do  not  form  any  distinct 
radial  rows  nor  any  very  evident  reticulations.  Most  of  those  on 
the  basal  part  of  the  rays  stand  in  short  transverse  or  oblique  rows  or 
combs,  varying  from  two  or  three  to  ten  or  more,  but  many  stand 


SHALLOW-WATER   STARFISHES  l6/ 

singly  or  in  small  clusters.  On  the  distal  third  of  the  rays  they 
become  much  more  crowded  and  stand  in  small  groups  or  singly; 
on  the  disk  they  form  irregular  short  rows  or  imperfect  reticulations. 
Everywhere  on  the  disk  and  basal  half  of  rays  there  are  large  papular 
areas,  with  very  large  groups  of  small  papulae.  The  spines  are  sur- 
rounded and  partly  concealed  by  small  wreaths  of  minute  minor 
pedicellariae,  which  become  more  abundant  on  the  distal  part  of  the 
rays.  Many  others  occur  among  the  papulae.  Major  pedicellariae 
occur  very  rarely  on  the  dorsal  surface  in  this  specimen,  and  only 
of  small  size.  They  are  much  more  frequent  on  the  submarginal 
channels  and  actinal  interbrachial  naked  areas ;  these  are  long-ovoid, 
not  very  large,  tapered,  and  sometimes  with  denticulate  tips.  Others 
of  small  size,  acute-ovate  in  form,  occur  within  the  margins  of  the 
ambulacral  furrows.  The  madreporic  plate  is  large,  very  porous, 
with  close,  very  complex,  narrow  gyri. 

The  dorsal  ossicles  are  very  numerous,  rather  slender,  and  loosely 
reticulated,  so  as  to  leave  very  large  papular  areas.  The  actinal 
ossicles  are  much  stronger  and  more  closely  united ;  and  though  nar- 
row they  are  rather  thick.  Seen  from  the  inside,  the  spaces  between 
them  are  deep,  squarish  and  pretty  regular.  The  ambulacral  ossicles 
are  very  numerous  and  much  compressed,  so  that  they  are  thin 
between  the  pores;  but  they  are  elongated  in  the  direction  of  the 
depth. 

The  type  and  only  large  specimen  of  this  species  was  taken  at 
Popof  Island,  Alaska,  by  the  Harriman  Expedition,  July  10,  1899, 
and  was  sent  to  me  by  Professor  Ritter.  It  was  unfortunately  dried 
with  the  rays  badly  bent,  and  some  of  them  were  broken  in  trans- 
portation. 

This  species  does  not  appear  to  be  very  closely  allied  to  any 
other,  except  E.  troschelii.  The  latter  has  more  slender  rays,  with 
the  dorsal  spines  distinctly  reticulated  and  strongly  capitate,  fewer 
rows  of  ventral  spines,  and  different  pedicellariae.  A  large  series  of 
specimens  might,  however,  show  that  it  is  only  an  overgrown  speci- 
men of  that  species,  or  of  its  variety  rudis. 

In  the  character  of  the  dorsal  spines,  it  resembles  Pisaster  con- 
fertus;  but  the  latter  is  strictly  monacanthid,  and  has  shorter  rays, 
a  much  broader  disk,  more  regularly  reticulated  dorsal  spines,  and 
different  pedicellariae. 


1 68  VERRILL 

Genus  Orthasterias  Verrill,  nov. 

Long-rayed  diplacanthid  starfishes  with  a  small  disk.  Dorsal 
skeleton  consists  of  stout,  wide,  subimbricated,  four  or  five-lobed 
plates  arranged  in  three,  five,  or  more  definite  longitudinal  radial 
rows,  and  sometimes  with  extra  interpolated  flat  ossicles,  the  rows 
connected  by  transverse  ossicles.  The  principal  plates  bear,  on  a 
central  boss,  large,  isolated  spines,  standing  in  three  to  five  or 
more  longitudinal  rows.  The  transverse  ossicles  are  strong,  oblong- 
elliptical. 

Usually  one  row  of  interactinal  or  peractinal  plates,  with  or  with- 
out spines,  closely  united  to  the  adambulacrals  and  inferomarginals ; 
sometimes  they  are  rudimentary,  and  without  spines;  rarely  two 
rows. 

The  inferomarginals  are  strong,  four-lobed  plates,  each  bearing 
usually  two  large  spines.  Superomarginals  are  large,  usually 
monacanthid,  four-lobed;  the  descending  lobe  is  large  and  usually 
joined  directly  to  the  inferomarginals,  by  overlapping  the  upper  lobe 
of  the  latter.  The  alternate  supermarginal  plates  are  usually  with- 
out a  spine  in  all  of  our  species  except  O.  forreri,  O.  leptolena,  and 
perhaps  O.  koehleri. 

Seen  from  the  interior,  the  dorsal  skeleton  is  largely  composed  of 
the  stout  connective  ossicles,  mostly  in  transverse  positions,  and  some- 
what reticulated. 

This  genus  is  represented  on  the  Atlantic  coast  of  the  United 
States  by  O.  tanneri  Ver.  and  by  0.  subangulosa  Ver.  (=Asterias 
angulosa  Per.,  name  preoccupied)  in  the  West  Indies.  O.  eustyla 
(Sladen)  seems  also  to  belong  to  the  genus. 

The  lateral  and  dorsal  major  pedicellariae  are  unusually  large,  and 
are  usually  of  two  or  more  forms.  The  larger  are  erect,  wedge- 
shaped,  stone-hammer-shaped,  or  ovate,  with  the  tips  of  the  valves 
wide  and  usually  denticulate.  Others  are  more  slender,  with  the 
valves  spatulate  or  narrowed  in  the  middle  and  the  tips  unguiculate, 
with  interlocking  teeth.  Others  similar,  or  smaller,  with  acute  tips, 
often  occur  on  the  adambulacral  spines  and  margin.  Type, 
O.  columbiana  V. 

ORTHASTERIAS  COLUMBIANA  Verrill,  sp.  nov. 

Plate  xxjv,  figures  i,  2,  4;  plate  xxxv,  figure  ij  plate  txv,  figure  2;  plate 
-j|f  LXXVIH,  figures  i-ie  (pedicellariae)  ;  plate  LJfxtrx,  figures  3-3*:  (spines  and 
pedicellariae)  ;  plate.ax,  figures  2-2&  (structure). 

The  type  of  this  species  is  a  rather  large  specimen  collected  at 
Yakutat,  Alaska,  by  Prof.  W.  R.  Coe,  on  the  Harriman  Expedition. 


SHALLOW-WATER   STARFISHES  169 

The  radii  are  18.5  mm.  and  157  mm. ;  ratio,  about  1 : 8.5.  The  disk 
is  small,  the  five  arms  are  long,  rounded  or  subangular  above,  and 
taper  very  gradually  to  rather  small  tips.  The  dorsal  plates  are 
numerous,  forming  five  rows,  with  some  interpolated  ones.  They 
are  large,  wide,  considerably  thickened  or  convex,  those  in  the  three 
primary  radial  rows  having  more  convex  centers  and  forming  a 
low  but  distinct  carina. 

The  dorsal  spines  form  about  five  more  or  less  regular  rows  above 
the  upper  marginals,  often  with  some  intermediate  ones.  The  five 
main  rows  extend  nearly  to  the  tips  of  the  rays,  the  spines  becoming 
rather  crowded  distally,  while  proximally  they  stand  well  apart, 
except  in  the  median  row.  The  dorsal  spines  are  all  nearly  equal, 
except  distally,  rather  large,  cylindrical,  or  slightly  clavate,  with 
obtuse  or  rounded  sulcate  tips,  and  bearing  dense  wreaths  of  minor 
pedicellariae  above  or  about  the  middle. 

The  larger  spines  are  about  4  mm.  high  and  I  mm.  in  diameter. 
The  upper  marginal  plates  are  also  thick  and  convex;  their  spines 
are  like  the  dorsals,  though  a  little  larger,  and  bear  similar  groups 
of  pedicellariae.  They  form  a  very  regular  row,  one  to  each  alternate 
plate.  Along  the  middle  of  the  ray  they  stand  about  3.5  mm.  to  4.5  mm. 
apart,  but  become  crowded  to  I  mm.  or  2  mm.  distally.  Between  the 
upper  and  lower  marginals  there  is  a  rather  wide  naked  channel, 
crossed  by  the  thickened  descending  apophyses  of  the  marginal  plates, 
and  bearing  on  the  plates  many  very  large,  ovate  and  spatulate  major 
pedicellariae.  These  often  alternate  with  the  spines;  others  are 
irregularly  placed. 

The  inferomarginal  spines  form  two  rows,  two  divergent  spines 
standing  obliquely  close  together  on  most  of  the  plates.  They  are 
similar  to  the  upper  ones  in  size  and  shape,  though  many  are  a  little 
compressed  and  more  strongly  sulcate  at  the  tips.  They  bear  large 
groups  of  minor  pedicellariae  on  their  upper  sides,  about  mid-height. 
The  inferomarginal  plates  are  strongly  convex,  with  the  outer  sur- 
face elliptical.  Another  regular  row  of  similar  spines  is  borne  by 
the  peractinal  plates,  close  to  the  adambulacrals,  each  corresponding 
to  about  five  pairs  of  the  latter.  They  extend  regularly  nearly  to 
the  tips  of  the  rays  and  have  papular  pores  between  them.  The  per- 
actinal plates  are  rounded  and  convex. 

The  adambulacral  spines  form  two  very  regular  and  equal  rows. 
They  are  slender,  scarcely  tapered,  terete,  or  slightly  compressed 
distally,  with  blunt  tips.  They  are  shorter  and  much  smaller  than  the 
adjacent  actinal  spines.  Those  on  the  adoral  plates  are  distinctly 


I7O  VERRILL 

longer  and  more  slender.  Four  contingent  pairs  of  plates,  besides 
the  two  apical  ones,  form  the  compressed  oral  carinse,  each  bearing 
a  single  spine.  The  epioral  spines  are  like  the  others.  The  perorals 
consist  of  an  apical  pair  of  shorter  and  stouter  spines  and  a  divergent 
auxiliary  spine,  of  about  half  their  length,  on  each  side. 

Major  pedicellariae  of  different  sizes  and  forms  are  scattered  over 
the  dorsal  surface  as  well  as  on  the  lateral  channels  and  on  the  mar- 
gins of  the  ambulacral  grooves;  others  occur  on  the  actinal  areas, 
on  the  tips  of  the  jaws,  and  on  the  adambulacral  spines.  The  largest 
occur  on  the  marginal  plates.  These  are  compressed,  long-ovate, 
oblong-ovate  or  oblong-lanceolate  in  a  front  view;  some  are  very 
stout,  with  somewhat  spatulate  rounded  tips,  in  a  profile  view ;  a  few 
are  denticulate,  others  acute.  Those  of  the  ambulacral  region  are 
more  compressed  and  usually  much  smaller,  though  the  larger  ones 
are  here  as  wide  as  the  adambulacral  spines ;  most  of  these  are  long- 
ovate  or  lanceolate  and  acute.  Those  on  the  back  are  mostly  acute- 
ovate. 

The  minor  pedicellariae  are  rather  unusually  large  and  form  dense 
clusters  around  all  the  dorsal  and  upper  marginal  spines  at  about 
mid-height,  while  smaller  groups  occur  on  the  actinal  and  adambu- 
lacral spines.  Papulae  are  small  and  numerous.  The  ambulacral 
grooves  are  wide;  the  pores,  which  form  four  regular  rows,  are 
narrow-elliptical,  with  flaring  lips. 

A  smaller  specimen  from  Queen  Charlotte  Islands  (Canadian  Geo- 
logical Survey,  1878)  has  the  radii  12  mm.  and  72  mm.;  ratio, 
i :  6.  The  dorsal  spines  are  more  numerous  and  longer  than  in  the 
type,  and  stand  in  about  five  rows.  They  are  all  nearly  equal  and 
rather  isolated,  except  those  in  the  median  radial  row,  where  they 
stand  in  a  pretty  dose  line.  They  all  bear  a  dense  wreath  of  minor 
pedicellariae  on  a  sheath,  at  about  mid-height.  The  dorsal  ossicles 
are  strong  and  closely  united,  convex,  with  a  mammilliform  elevation 
and  a  central  pit,  where  the  spine  was  attached. 

The  upper  and  lower  marginal  spines  are  a  little  longer,  but  similar 
to  the  dorsals ;  both  are  in  regular  rows,  but  the  lower  ones  often 
stand  two  on  a  plate  proximally.  Actinal  spines  similar,  in  one  row. 
Adambulacral  spines  long  and  slender,  but  rather  shorter  than  the 
actinals,  except  on  the  adoral  and  epioral  plates,  where  they  become 
decidedly  longer.  They  stand  irregularly,  one  and  two  to  a  plate. 
Adoral  carinae  compressed,  formed  by  two  or  three  contingent  plates, 
besides  the  epiorals.  Peroral  spines  of  moderate  length. 


SHALLOW-WATER   STARFISHES  171 

A  good  dry  specimen  taken  at  Departure  Bay,  British  Columbia, 
in  18  fathoms,  gravel  bottom,  September,  1908  (C.  H.  Young),  has 
been  sent  to  me  by  the  Canadian  Geological  Survey. 

Radii,  28  mm.  and  190  mm. ;  ratio,  about  i :  7.  The  dorsal  skeleton 
is  rather  firm,  though  the  plates  are  small,  but  the  plates  and  cross- 
bars are  numerous,  and  papular  areas  are  not  very  large.  The 
median  row  of  spines  is  pretty  close  and  regular,  composed  of  a 
single  row  of  large  and  long,  nearly  cylindrical,  partly  tapered,  blunt 
spines,  one  to  a  plate.  On  each  side  there  are  two  or  three  less 
regular,  rather  distinct  rows  of  widely  spaced  long  spines,  like  those 
of  the  median  row,  but  not  half  so  numerous. 

Below  the  middle  of  the  side  there  is  a  more  regular  row  of  similar 
but  more  numerous  superomarginal  spines,  curving  upward,  proxi- 
mally,  to  the  upper  margin  of  the  interradial  angles.  These,  like  the 
dorsal  spines,  have  a  close  wreath  of  small  minor  pedicellariae, 
usually  near  the  tips,  but  at  least  beyond  the  middle. 

Separated  from  the  upper  marginal  plates  by  a  wide,  naked, 
papular  lane,  there  is  a  regular  double  row  of  large  spines,  which 
are  the  inferomarginals.  These  spines  are  shorter  and  rather 
smaller  than  those  above,  with  blunt,  fluted  tips.  They  stand 
obliquely  on  the  plates,  the  lower  one  more  distal,  and  close  to  a 
similar  spine  arising  from  each  of  a  series  of  small,  roundish,  per- 
actinal  plates,  so  that  the  three  rows  of  spines  form  a  series  of 
obliquely  transverse,  short,  close  rows,  only  slightly  separated  from 
the  adambulacrals.  Each  peractinal  plate  corresponds  to  four  or  five 
adambulacrals. 

The  adambulacral  spines  are  arranged  regularly,  two  to  a  plate. 
They  are  equal,  slender,  tapered,  subacute,  shorter  and  much  smaller 
than  the  adjacent  ventral  spines.  Carried  on  the  adambulacral  spines 
and  within  the  borders  of  the  grooves,  are  numerous  rather  large, 
lanceolate,  subacute  major  pedicellariae,  many  of  them  equal  in 
thickness  to  that  of  the  spines,  at  mid-height,  and  from  one-fourth 
to  one-third  as  long.  On  the  sides  and  back  of  the  rays  there  are 
also  many  scattered  major  pedicellarise,  mostly  of  still  greater  size. 
Some  of  these  are  lanceolate,  but  most  are  thick,  stout,  ovate  or 
oblong-ovate,  with  the  tips  of  the  jaws  blunt  and  strongly  denticu- 
late. 

The  dried  specimen  has  been  stained  by  the  preparator  to  a  deep 
red-brown  color,  probably  imitating  its  color  in  life. 

The  largest  specimen  that  I  have  seen  is  about  two  feet  in  diameter, 
as  dried,  it  has  the  greater  radii  340  mm.;  lesser,  30  mm.;  ratio, 


172  VERRILL 

1 : 8.  It  was  taken  in  25  fathoms,  off  Victoria,  British  Columbia,  and 
was  sent  to  me  by  Dr.  C.  F.  Newcombe,  from  the  Provincial 
Museum  of  British  Columbia. 

It  agrees  well  with  the  specimen  from  Departure  Bay,  in  nearly 
all  characters,  except  those  apparently  due  to  greater  age.  The  five 
rows  of  large  dorsal  spines  are  pretty  regular  and  are  arranged  as  in 
the  latter,  as  are  the  marginal  and  ventral  spines.  As  in  the  smaller 
specimen,  there  are  but  three  rows  of  ventral  spines,  standing  in 
the  same  way,  in  short  oblique  groups  of  three,  but  these  spines  are 
relatively  longer,  being  nearly  as  long  as  the  upper  marginals. 
Major  pedicellariae  are  of  different  sizes  and  forms,  but  mostly  large, 
scattered  on  the  back  and  sides,  as  already  described.  The  large 
denticulate  form  is  most  numerous. 

Alcoholic  specimens  of  this  species  were  also  obtained  by  the  Har- 
riman  Expedition  (pi.  xxrv,  figs.  I,  2).  These  have  the  rays  plump 
and  rounded.  The  wreaths  of  minor  pedicellariae  on  the  dorsal  and 
lateral  spines  are  so  large  that  they  are  nearly  or  quite  in  contact, 
over  much  of  the  surface.  The  ambulacral  feet  are  large  and 
crowded.  The  spines  are  arranged  nearly  as  in  the  dry  specimen 
described  above. 

This  species  is  evidently  closely  allied  to  0.  dawsoni,  which  I  for- 
merly supposed  to  be  merely  the  young.  But  the  latter  has  much 
fewer  dorsal  and  lateral  spines  and  plates,  owing  to  the  relatively 
larger  size  of  the  latter.  This  is  especially  noticeable  when  the  distal 
parts  of  a  ray  of  each  are  compared,  where  of  the  same  size ;  for  the 
present  species  has  at  least  twice  as  many  plates  and  spines  in  the 
same  space.  The  forms  of  the  spines  and  pedicellariae  are  somewhat 
different,  though  similar.  The  arrangement  of  the  adambulacral 
and  interactinal  spines  is  nearly  the  same  in  both,  when  of  similar 
size.  The  larger,  lateral,  spatulate  pedicellariae  are  more  spatulate 
or  lyrate  and  longer  in  dawsoni,  but  I  have  found,  in  the  latter,  none 
of  them  serrate  or  unguiculate,  while  in  this  some  occur  with  three 
or  four  small  interlocking  apical  teeth. 

This  species  is  remarkable  for  its  thick  and  enlarged  dorsal 
plates,  which  are  united  so  closely  together  that  the  papular  areas 
are  mostly  small.  The  small  interpolated  ossicles  are  flat  and  angular. 
The  marginal  and  interactinal  plates  are  also  unusually  thick  and 
convex. 

Yakutat,  Alaska  (Prof.  W.  R.  Coe,  Harriman  Expedition) ; 
Queen  Charlotte  Islands  (Canadian  Geological  Survey,  1878) ; 
Departure  Bay,  British  Columbia,  18  fathoms  (C.  H.  Young,  Cana- 


SHALLOW- WATER   STARFISHES  173 

dian  Geological  Survey) ;  off  Victoria,  British  Columbia,  in  25  fath- 
oms (C.  F.  Newcombe,  Prov.  Mus.  B.  C),  very  large. 

ORTHASTERIAS  BIORDINATA  Verrill,  sp.  nov. 

Plate  Lxm,  figures  i,  2  (general  of  type)  ;  plate  LXXXII,  figures  2-26  (spines 

and  pedicellariae). 

The  two  type-specimens  are  very  much  alike  and  nearly  of  the 
same  size.  Radii  of  the  one  figured,  8  mm.  and  88  mm.;  ratio, 
i :  ii ;  radii  of  the  other,  9  mm.  and  85  mm. ;  ratio,  1:9:5. 

Rays  five,  long,  slender,  very  gradually  tapered,  rather  acute,  well 
rounded  above,  and  furnished  with  three  regular  rows  of  well 
spaced,  elongated  spines,  one  to  a  plate,  except  near  the  ends  of  the 
rays,  where  there  are  five  rows,  and  the  spines  are  closer  together. 
Isolated  spines  also  stand  on  the  transverse  ossicles,  here  and  there. 
The  dorsal  spines  are  regularly  clavate,  evenly  fluted  distally,  with 
the  tips  obtusely  rounded. 

They  stand  on  elevations  of  stout,  lobed  plates.  The  median 
row  is  distinct,  because  the  spines  are  about  twice  as  many,  though 
of  about  the  same  size  and  length.  The  center  of  the  disk  is  occu- 
pied by  a  prominent  five-lobed  plate,  bearing  a  spine,  and  with  five 
stout  ossicles  radiating  out  from  its  lobes,  in  line  with  the  median 
rows  of  the  rays,  but  connecting  with  five  radial  spine-bearing 
plates  of  the  disk,  which  form  a  pentagon. 

Alternating  with  these,  and  a  little  farther  away,  are  five  larger 
interradial  plates,  each  bearing  a  spine  of  larger  size. 

The  dorsal  pore  is  small  and  surrounded  by  minute  papillae.  It  lies 
between  two  lobes  of  the  central  plate  and  in  an  interradius  next  to 
that  occupied  by  the  madreporite,  which  is  large,  very  round,  and 
with  fine  gyri. 

The  spines  are  surrounded  by  close  wreaths  of  very  small  minor 
pedicellarige  which  are  often  subbasal,  but  frequently  at  mid-height, 
or  distal,  and  then  are  attached  to  the  edges  of  a  sheath. 

The  superomarginals  form  a  very  regular  row,  one  to  a  plate,  of 
rather  longer  and  more  slender  fluted  spines.  The  inferomarginal 
row  is  double,  with  two  spines  to  a  plate;  these  spines  are  dis- 
tinctly longer,  more  slender,  and  less  clavate  than  the  dorsals,  but 
fluted  in  the  same  way.  Alternate  superomarginals  are  spineless. 

The  peractinals  are  smaller;  on  several  rays  they  extend  only  on 
the  proximal  third  of  the  ray ;  on  others  to  the  distal  fourth. 

The  adambulacral  spines  are  slender,  regularly  diplacanthid, 
widely  divergent,  forming  two  remarkably  regular  pectinate  rows, 


174  VERRILL 

the  inner  one  with  the  spines  horizontal  and  meeting  or  interlocking 
over  the  middle  of  the  groove  in  most  places. 

The  dorsal  dermal  major  pedicellariae  are  large,  long,  mostly 
compressed,  with  spatulate  and  denticulate  valves;  those  between 
the  marginal  spines  are  larger,  with  oblong  or  spatulate  valves, 
obtuse  and  dentate  at  the  tips  (see  pi.  LXXXII,  figs.  2-26).  Colors 
yellowish  brown,  the  disk  reddish  brown. 

The  type  specimens  are  from  Departure  Bay,  British  Columbia 
(coll.  C.  M.  Young,  Canadian  Geological  Survey). 

This  may  be  merely  a  variety  of  the  young  of  O.  columbiana, 
coming  from  the  same  district,  but  I  have  seen  no  intermediate 
specimens.  The  remarkable  regularity  of  the  adambulacral  spines 
is  a  striking  feature,  as  well  as  the  regular  spaced  rows  of  dorsal 
and  marginal  spines. 

ORTHASTERIAS  CALIFORNICA  Verrill,  sp.  nov. 

LO  H  $ 

Plate  LXVIII,  figure  2  (actinal  side  of  type)  ;  plate  LXX,  figure  5  (actinal  side)  ; 
plate  LXXX,  figures  3-30  (spines  and  pedicellariae)  ;  plate  LXXXI,  figures 
2-2&  (pedicellariae  and  spines)  ;  text-figure  3.  «  ,$"? 

Rays  five,  elongated,  narrow,  well  rounded  above,  constricted  at 
base.  Disk  small.  Radii,  9  mm.  and  86  mm. ;  ratio,  1 : 9.5. 

Dorsal  and  marginal  skeletal  ossicles  strong,  arranged  in  rather 
irregular  longitudinal  rows,  each  of  the  dorsals  and  superomarginals 
bearing  a  single,  rather  elongated,  stout,  cylindric  or  little  tapered, 
blunt  spine.  The  spines  are  pretty  evenly  spaced,  nearly  equal  and 
surrounded  above  the  base  (as  dried)  by  a  dense  wreath  of  small, 
ovate  minor  pedicellariae.  The  .abactinal  spines  form  five  indistinct 
rows  proximally,  but  only  three  on  the  distal  fourth  of  the  rays. 
They  are  remarkably  uniform  in  size  and  length.  The  median  row 
is  scarcely  different  from  the  others.  Those  on  the  disk  are  similar 
and  regularly  spaced. 

The  superomarginals  form  a  single,  regular  row ;  they  are  like  the 
dorsals,  but  rather  longer.  The  inferomarginals  are  decidedly 
smaller  and  somewhat  shorter.  They  stand  mostly  two  to  a  plate; 
the  upper  one  of  the  pair  bears  a  half-wreath  of  minor  pedicellariae. 
These  spines  are  mostly  flattened  at  the  tips.  On  the  proximal  half 
of  the  rays  there  is  a  simple  row  of  peractinal  spines,  similar  to  the 
adjacent  marginals  and  close  to  them. 

Adambulacral  spines  are  mostly  two  to  a  plate,  rather  long,  very 
slender,  flattened,  larger  toward  the  mouth.  Each  jaw  has  a  terminal 
pair  of  short,  notably  stout,  blunt,  peroral  spines,  and  a  much  longer 
pair  of  tapered  epiorals. 


SHALLOW- WATER   STARFISHES  175 

Sessile  major  pedicellariae  are  large,  erect.  The  larger  nearly 
equal  to  the  spines  in  diameter,  are  sparsely  scattered  on  the  dorsal 
side  and  more  frequently  on  the  intermarginal  channels;  some 
equally  large  are  also  attached  to  the  inner  edges  of  the  ambulacral 
grooves,  and  in  some  of  the  interradial  areas  below,  but  most  of  the 
axils  bear  groups  of  much  smaller  ones.  The  larger  pedicellariae  are 
somewhat  elongated  wedge-shaped,  with  broad,  spatulate  jaws, 
strongly  dentate  or  unguiculate  at  the  wide  tips. 

The  madreporite  is  large,  flat,  fine-grained,  flush  with  the  surface, 
with  a  few  spines  and  large  pedicellarise  around  it,  but  with  no 
special  bordering.  Ocular  or  apical  plates  covered  with  many  small 
spines. 

Off  San  Francisco,  probably  in  rather  shallow  water  (Prof.  W.  E. 
Ritter). 

ORTHASTERIAS  KCEHLERI  (de  Loriol). 

Plate  LXXV,  figures  3-3^  (type,  after  de  Loriol). 

Asterias  kcehleri  DE  LORIOL,  Mem.  Soc.  Phys.  et  d'Hist.  Nat.  Geneve,  xxxn, 
No.  9,  p.  21,  pi.  m  (xvm),  figs.  3-5,  1897. 

The  type  had  the  following  characters : 

Rays  five,  long,  slender,  well  rounded  dorsally.  Radii,  10  mm.  and 
97  mm. ;  ratio,  i :  9.7.  Dorsal  spines  numerous,  long  (2  mm.  to 
2.5  mm.),  slender,  blunt,  striated  distally,  arranged  without  order. 
Ventral  spines  [marginal  and  peractinal]  form  three  regular  rows ; 
they  are  rather  larger  and  longer  than  the  dorsals.  Adambulacral 
spines  long  and  slender,  mostly  two  to  a  plate,  divergent. 

A  few  large,  dermal,  dorsal  pedicellariae,  some  of  them  I  mm. 
long;  they  are  long-lanceolate,  acute,  dentate  at  the  tip.  Major 
pedicellariae  of  unequal  sizes  occur  on  the  actinal  side  and  on  the 
edges  of  the  ambulacral  grooves. 

Saanich  Inlet,  Vancouver  Island  (de  Loriol). 

ORTHASTERIAS  DAWSONI  Verrill,  sp.  nov. 

Plate  xxm,  figures  i,  2  (type)  ;  plate  LXXV,  figures  2-26  (spines  and  pedicel- 
lariae) ;  plate  LXXX,  figures  2,  a-g  (pedicellariae)  ;  plate  LXXXI,  figures 
3~3&  (spines  and  pedicellariae). 

Disk  very  small;  rays  five,  slender,  angular  above  when  dry, 
gradually  tapered,  Radii  of  type,  10  mm.  and  80  mm. ;  ratios,  i :  8. 

Dorsal  spines  in  three  regular  radial  rows,  long,  tapered,  obtuse, 
or  subacute,  standing  singly  on  the  plates. 


<*&#  2.  t 


176  VERRILL 

Superomarginal  spines  similar  to  dorsals,  in  a  simple  regular  row. 
Each  alternate  plate  usually  bears  a  large  spine.  On  the  disk  about 
fifteen  similar  spines  form  a  circle  around  the  center,  a  few  spines 
standing  within  it.  The  circle  is  formed  by  five  pairs  of  interradial 
spines  and  five  isolated  radials. 

The  stout  dorsal  ossicles  are  articulated  transversely  and  longi- 
tudinally, so  as  to  leave  large,  squarish  interspaces,  carrying  numer- 
ous small  papulae.  Minor  pedicellariae  form  dense  wreaths  around  all 
the  dorsal  and  upper  marginal  spines  at  about  mid-height. 

Madreporic  plate  small,  flat,  with  fine  gyri.  The  upper  marginal 
plates  are  stout,  with  a  broad,  obtuse  apophysis  extending  down  to 
the  lower  marginals,  but  leaving  triangular  papular  areas  between, 
so  that  there  is  a  wide  naked  lane  between  the  upper  and  lower  mar- 
ginals. A  few  very  large  major  pedicellariae,  with  wide  spatulate 
blades,  occur  on  this  area.  The  lower  marginal  plates  are  also  stout 
and  prominent,  each  one  usually  bearing  two  prominent,  subacute, 
divergent  spines,  which  are  nearly  as  long  as  the  upper  ones,  but 
much  more  slender  and  twice  as  numerous.  Their  large  clusters  of 
pedicellariae  are  mostly  confined  to  their  upper  side.  A  less  regular 
row  of  similar  spines  is  borne  on  the  small  peractinal  plates,  each  of 
which  corresponds  to  about  three  adambulacrals. 

The  adambulacral  spines  form  two  very  regular  rows  of  rather 
long,  slender,  slightly  tapered  spines,  which  are  often  flattened  at 
the  tips.  Those  of  the  inner  row  are  a  little  shorter  than  the  outer 
ones,  and  more  acute.  About  four  of  the  adambulacral  plates  cor- 
respond with  one  marginal.  There  is  a  row  of  small  papular  areas 
between  the  peractinals.  A  few  small  ovate  pedicellariae  sometimes 
occur  on  the  adambulacral  spines  and  in  the  grooves.  The  small 
actinal  interradial  areas  bear  groups  of  small,  long-ovate  major  pedi- 
cellariae of  several  sizes. 

The  very  large  dermal  major  pedicellariae  on  the  sides  of  the  rays 
are  not  compressed,  often  nearly  as  stout  as  the  adjacent  spines,  and 
in  a  profile  view  the  blades  are  broadly  spatulate  with  abruptly 
rounded  ends,  but  not  unguiculate  nor  distinctly  serrulate,  so  far 
as  observed. 

Ambulacral  grooves  wide ;  sucker-feet  large,  placed  in  four  regu- 
lar rows. 

Queen  Charlotte  Islands  (Dr.  G.  M.  Dawson,  Canadian  Geological 
Survey,  1875). 


SHALLOW- WATER   STARFISHES  177 

ORTHASTERIAS  MERRIAMI  Verrill,  sp.  nov. 

Plate  xvm,  figures  4,  5  (type)  ;  plate  xix,  figures  i,  2;  plate  LXXV,  figures  l-id 

(spines  and  pedicellariae,  No.  1181).  r  k.«xUvi/vv*HJl 
A  large  six-rayed  species,  with  long,  round,  gradually  tapered 
rays.     Disk  of  moderate   size.     In   the  type-specimen,   from  off 
Juneau,  Alaska,  the  radii  are  17  mm.  and  112  mm.;  ratio,  about 

1:6.5. 

Dorsal  ossicles  broad,  strong,  convex,  and  firmly  united,  with 
small  papular  spaces  intervening;  spines  well  spaced,  conspicuous, 
stout,  but  not  very  long,  cylindrical  or  a  little  tapered,  obtuse.  They 
stand  singly  on  the  ossicles  and  are  surrounded  at  base  by  large 
wreaths  of  pedicellariae.  They  form  an  irregular  radial  row  and 
two  or  three  indistinct  or  imperfect  rows  each  side  of  it,  with 
smaller  spines  interpolated.  The  upper  and  lower  marginal  spines 
form  equal  regular  rows,  one  spine  on  each  plate.  These  spines  are 
larger  and  longer  than  the  dorsals,  mostly  conical  and  subacute ;  those 
in  the  lower  series  are  distinctly  longer,  especially  near  the  base 
of  the  rays.  Those  in  the  upper  row  bear  large,  dense,  complete 
wreaths  of  pedicellariae;  those  in  the  lower  row  have  them  only  on 
the  upper  side.  The  two  rows  are  separated  by  a  naked  channel 
which  becomes  wide  proximally  and  bears  large,  ovate  major  pedi- 
cellariae, clusters  of  minor  pedicellariae,  and  a  row  of  papular  pores. 
At  the  base  of  the  rays  are  two  close  rows  of  interactinal  spines, 
similar  to  the  lower  marginals.  Of  these,  the  outer  row  is  close  to 
the  marginal,  but  with  small  intervening  papular  pores,  and  it 
extends  nearly  to  the  end  of  the  ray.  Its  ossicles  are  rounded,  con- 
vex, and  nearly  as  large  as  the  marginals.  The  inner  row  extends 
only  about  half  the  length  of  the  ray.  Many  of  the  actinal  spines 
are  flattened  or  acuminate  at  the  tip ;  others  are  obtuse. 

The  adambulacral  spines  on  the  middle  and  distal  plates  stand  one 
or  two  to  a  plate,  alternating  irregularly,  but  on  the  proximal  fourth 
of  the  ray  they  are  mostly  one  to  a  plate.  They  are  rather  stout, 
shaped  like  the  actinal  spines,  but  smaller  and  shorter,  varying  in 
size,  mostly  obtuse,  but  the  smaller  ones  often  acute.  Toward  the 
mouth  they  become  longer,  more  slender  and  subacute.  The  peroral 
spines  are  large  and  strong  and  nearly  meet  over  the  mouth.  The 
two  apical  peroral  spines,  which  are  much  stouter  than  the  adorals, 
but  not  so  long,  are  straight,  tapered,  a  little  flattened,  obtuse ;  the 
smaller  side-spine  is  about  half  as  long,  but  of  the  same  shape. 
They  bear  small,  ovate,  forficulate  pedicellarise.  The  epiorals  and 
adorals  are  similar  in  form,  distinctly  longer  than  those  farther  out, 
13 


178  VERRILL 

slender,  terete,  and  evenly  tapered.  The  adoral  carina  consists  of 
three  pairs  of  closely  contingent  plates,  besides  the  epiorals.  Large, 
ovate,  major  pedicellariae  occur  between  the  interactinal  spines. 

The  major  pedicellariae,  between  and  above  the  marginal  spines, 
are  still  larger,  strongly  compressed,  ovate,  obtuse,  not  serrate.  The 
minor  pedicellariae  not  only  form  large  wreaths  around  the  middle  of 
the  dorsal  and  lateral  spines,  but  also  occur  in  clusters  on  the  papu- 
lar areas. 

A  smaller  specimen  (pi.  xix,  figs.  I,  2)  was  taken  at  nearly  the 
same  place  as  the  type,  and  is  now  preserved  in  alcohol.  According 
to  Dr.  Coe's  notes  the  color  of  this,  in  life,  was  nearly  uniform  deep 
yellow,  but  paler  ochre-yellow  on  the  back.  Radii,  14  mm.  and 
66  mm. ;  ratio,  about  1 : 4.7.  The  larger  dorsal  spines  are  in  five 
irregular  rows,  with  some  smaller  ones  interpolated.  Only  a  few  of 
the  proximal  adambulacral  plates  are  monacanthid. 

In  alcohol  it  has  a  rigid  appearance  and  is  firm  to  the  touch,  owing 
to  the  abundant  flat  dorsal  ossicles.  The  principal  spines  are  con- 
spicuous above  the  general  surface,  blunt  and  sulcated  at  the  tips,  but 
are  surrounded  proximally  by  thick,  fleshy  sheaths.  The  wreaths  of 
minor  pedicellariae,  which  are  attached  to  the  sheaths,  are  regular 
and  dense,  and  with  the  papulae  they  entirely  and  closely  cover  the 
intervals  between  the  spines.  The  larger  ventral  spines  are  conical, 
acute,  and  stand  in  two  very  regular  rows.  A  few  smaller  spines 
belong  to  a  synactinal  series.  The  ambulacral  feet  are  large  and 
have  large  suckers. 

This  species  is  known  only  from  a  few  specimens.  One  was  taken 
off  Juneau,  in  20  fathoms,  June  6;  another,  in  Glacier  Bay,  10 
fathoms,  June  10,  by  the  Harriman  Expedition,  1899  (W.  R.  Coe). 
Another  is  from  the  Gulf  of  Georgia  (No.  1181,  Mus.  Comp.  Zool.). 
The  species  is  dedicated  to  Dr.  C.  Hart  Merriam. 

The  only  west  American  species  having  any  marked  resemblance 
to  this,  when  adult,  is  O.  columbiana.  The  latter,  however,  is  a  five- 
rayed  species,  with  the  dorsal  spines  fewer,  much  longer,  and  in  more 
definite  rows.  Its  marginal  and  actinal  spines  are  also  longer  and 
more  cylindrical,  but  they  are  arranged  nearly  in  the  same  way.  Its 
adambulacral  spines  are  much  more  slender  and  form  two  regular 
rows.  Its  major  pedicellariae  are  larger,  stouter,  and  differently 
shaped.  When  very  young  this  species  probably  resembles  Lept- 
asterias  coei,  or  some  of  the  allied  species  with  six  rays,  but  would 
probably  have  fewer  marginal  and  actinal  spines. 


SHALLOW- WATER   STARFISHES  179 

Subgenus  Stylasterias  Verrill,  nov. 

Interactinal  plates  rudimentary  or  small,  disk-like,  standing  edge- 
wise, and  spineless.  See  p.  48. 

ORTHASTERIAS  FORRERI  (de  Loriol). 

Plate  LXV,  figure  i  (dorsal  side)  ;  plate  LXVI,  figures  i,  2  (dorsal  and  actinal 
sides)  ;  plate  LXX,  figure  7  (portion  enlarged)  ;  plate  LXXVII,  figures  i-id 
(spines  and  pedicellariae)  ;  plate  LXXX,  figures  i-ie  (pedicellarise,  etc.). 

Asterias  forreri  DE  LORIOL,  Notes  pour  servir  a  1'etude  des  Echinodermes, 
No.  II,  Recueil  zoologique  Suisse,  iv,  No.  3,  June  23,  p.  401,  pi.  18,  fig. 
i,  i88jf.  7  Jennings,  Johns  Hopkins  Univ.  Circular,  No.  195,  p.  16,  1907 
(habits,  name  not  used)  ;  Univ.  Calif.  Publ.,  Zool.,  iv,  pp.  53-185,  19  text- 
figures,  1907  (behavior). 

A  five-rayed  species  with  long,  gradually  tapered,  convex  rays, 
covered  with  long,  tapered,  well  spaced  spines  and  with  a  multitude 
of  remarkably  large,  unguiculate  minor  pedicellariae  in  large  cir- 
cumspinal  wreaths. 

The  largest  specimen  (No.  1823)  has  the  radii  20  mm.  and 
215  mm.;  ratio,  i:  10.75.  Another  from  the  same  place  has  them 
ii  mm.  and  116  mm. ;  ratio,  1 : 10.50.  A  third  specimen  (No.  1431) 
is  intermediate  in  size. 

The  dorsal  spines  stand  mostly  in  three  pretty  regular  rows,  one 
to  a  plate,  but  usually  with  an  imperfect,  more  lateral  row  on  each 
side  proximally.  They  are  elongated,  up  to  6  mm.,  tapered,  terete, 
obtuse  or  subacute,  nearly  smooth,  not  fluted,  surrounded  basally  by 
very  broad,  loose  wreaths  of  notably  large,  sharply  unguiculate  minor 
pedicellariae.  The  superomarginal  spines  are  similar,  rather  longer, 
one  to  a  plate,  in  a  regular  row,  rather  low  down  on  the  side. 

The  inferomarginals  are  still  larger,  up  to  8  mm.  on  the  larger 
specimen,  placed  two  to  a  plate,  pretty  near  the  adambulacrals.  They 
are  mostly  obtuse,  with  the  tips  wider  and  flattened,  or,  in  the  case  of 
the  lower  one,  often  gouge-shaped.  There  are  no  peractinal  spines. 

The  adambulacral  spines  are  two  to  a  plate,  close  together,  form- 
ing two  pretty  regular  close  rows.  They  are  slender,  unequal;  the 
outer  one  is  longer  and  larger,  usually  flattened  and  wider  at  the  tip, 
and  commonly  gouge-shaped. 

The  orals  are  shorter,  decidedly  stouter,  flattened,  somewhat  cla- 
vate  and  blunt. 

The  dorsal  and  lateral  papular  areas  are  large.  In  the  narrow 
lane  between  the  inferomarginal  and  adambulacral  spines  there  is  a 
row  of  small  papular  areas;  peractinal  plates  small,  flat,  standing 
edgewise,  seldom  visible  without  cleaning ;  easily  seen  from  inside. 


l8o  VERRILL 

The  major  pedicellariae  are  large  and  of  several  forms.  The 
larger  dorsal  ones  are  large,  stout,  erect,  wedge-shaped,  blunt,  with 
several  small  terminal  denticles  on  each  valve.  They  are  often  as 
thick  as  the  adjacent  spines.  Others  are  more  slender,  with  spatulate 
valves,  terminating  in  about  three  curved,  interlocking  teeth.  The 
large  marginal  ones  are  very  large,  ovate,  wedge-shaped  or  hammer- 
head-shaped, with  dentate  valves.  Those  on  the  adambulacral 
borders  and  spines  are  few  and  smaller.  The  madreporite  is  rather 
large,  with  very  fine  gyri,  without  special  bordering  spines. 

Monterey  Bay,  California,  Nos.  18230,  1823^  types;  No.  1431, 
Alaska  ?  (Mus.  Comp.  Zool.). 

This  species  is  easily  distinguished  from  all  those  related  to  it  by 
the  unusually  large  and  peculiar  abundant  minor  pedicellariae,  and 
from  most  large  species  by  the  small  number  of  rows  of  dorsal 
spines.  The  alternate  superomarginal  plates  are  not  spineless,  as  in 
several  allied  species. 

ORTHASTERIAS  FORRERI  FORCIPULATA  Verrill. 

Plate  LXII,  figures  2,  3  (dorsal  and  actinal  of  ray)  ;  plate  LXX,  figure  9;  plate 
LXXXVIII,  figures  6,  60  (pedicellarise). 

Asterias  (Urasterias)  forcipulata  VERRILL,  Amer.  Journ.  Sci.,  xxvui,  p.  67, 
1909;  Amer.  Naturalist,  XLVIII,  p.  542,  1909. 

Rays  long  and  slender,  gradually  tapered ;  length  of  ray,  325  mm. ; 
breadth,  not  including  spines,  28  mm. ;  disk  small.  Dorsal  skeleton 
weak,  with  large  papular  areas  nearly  concealed  by  vast  numbers  of 
unusually  large  minor  pedicellariae. 

The  dorsal  plates  are  small,  three-  or  five-lobed  or  stellate,  each 
of  the  larger  ones  usually  bearing  a  rather  long,  tapered,  subacute 
spine ;  these  are  well  spaced  and  form  about  five  irregular  or  indefi- 
nite rows.  The  spines  are  surrounded  by  wreaths  of  the  large  minor 
pedicellariae,  but  these  also  occur  in  larger  clusters  scattered  over  the 
integument  between  the  spines.  Large  major  pedicellariae  are  also 
scattered  over  the  back ;  these  are  stout,  ovate-lanceolate,  with  obtuse 
tips,  which  are  usually  strongly  denticulate. 

On  the  sides  of  the  ray  and  separated  from  those  above  by  a  wide 
papular  lane  there  is  a  row  of  small,  mostly  four-lobed  superomar- 
ginals,  usually  bearing  a  single  long  spine.  They  are  connected  to 
those  above  by  weak  transverse  ossicles  in  series,  leaving  large  papu- 
lar areas  between.  The  spines  are  rather  longer  and  larger  than  those 
of  the  dorsal  surface.  Between  these  and  the  adambulacral  spines 
there  is  a  single  row  of  stouter  spine-bearing  plates,  the  inferomar- 


SHALLOW-WATER   STARFISHES  l8l 

ginals ;  each  corresponds  to  five  or  six  adamibulacrals.  Most  of  these 
bear  two  long,  tapered  spines,  usually  blunt  and  somewhat  flattened  or 
sulcate  at  the  tips,  rather  larger  than  the  upper  marginals,  usually 
7  mm.  to  8  mm.  long.  Between  their  bases  there  are  often  scattered 
large  and  strong  denticulate  major  pedicellariae,  similar  to  those  of  the 
back,  but  mostly  stouter  and  more  obtuse ;  with  these  are  some  that 
are  much  smaller,  lanceolate,  and  subacute.  The  large  pedicellariae 
also  occur  on  the  naked  lanes  below,  both  on  the  papular  areas  and  on 
the  adambulacral  plates.  There  are  a  few  small,  thin,  synactinal 
ossicles  connecting  some  inferomarginals  to  the  adambulacrals,  but 
not  bearing  spines.  The  adambulacral  spines  form  two  regular  close 
rows,  two  on  each  plate;  they  are  slender,  tapered,  mostly  flattened, 
subacute,  about  5  mm.  to  5.5  mm.  long.  The  ambulacral  pores  are 
large  and  form  four  rows. 

The  dorsal  minor  pedicellariae  are  remarkable  for  their  great  size 
and  abundance ;  in  life  they  probably  nearly  conceal  the  whole  upper 
surface  and  spines,  and  are  borne  on  slender  pedicels.  Their  blades 
are  strongly  bent  and  very  strongly  dentate.  They  are  much  like 
those  of  O.  forreri. 

Their  jaws  are  slender,  strongly  arched,  open  widely  and  usually 
have  thin,  sharp,  spatulate  tips,  that  look  like  sharp  hooks,  when 
seen  in  profile.  They  are  about  i  mm.  in  height.  They  are  not  very 
unlike  those  on  U.  linckii,  but  much  larger.  Color  (when  dry),  gray- 
ish brown  above,  dull  yellow  below. 

The  type  was  from  Departure  Bay,  British  Columbia,  18  fathoms, 
gravel  (C.  H.  Young,  Canadian  Geological  Survey,  September, 
1908).  The  disk  was  not  sent  to  me.  Five  loose  rays  were  received. 

Although  this  subspecies  is  somewhat  allied  to  U.  linckii,  and  re- 
sembles it  in  form  and  the  arrangement  of  the  dorsal  spines,  it  differs 
widely  in  several  characters.  U.  linckii  has  more  numerous  and 
larger  sacculated  spines.  It  has  large  groups  and  wreaths  of  minor 
pedicellariae  of  unusually  large  size,  but  they  are  not  half  so  large  as 
in  this  species  and  are  far  less  abundant  dorsally,  the  larger  wreaths 
being  on  the  lateral  spines.  They  also  differ  in  form.  The  large, 
denticulate  major  pedicellariae  are  also  abundant  on  U.  linckii,  dor- 
sally, laterally,  and  in  the  grooves  (intra-ambulacral),  but  they  are 
not  half  so  large  in  this  species.  In  the  former  the  adambulacral 
plates  and  spines  are  much  less  numerous  and  less  crowded,  so  that 
only  about  four  of  the  plates  correspond  to  one  inferomarginal,  while 
in  this  species  there  are  six  or  seven  to  one  inferomarginal.  In 
U.  linckii  each  adambulacral,  on  the  proximal  half  or  more  of  the 


l82  VERRILL 

ray,  usually  bears  a  single  acute  spine,  though  on  the  distal  part  each 
bears  two  divergent  spines,  of  which  the  inner  is  much  smaller  and 
shorter.  But  in  the  present  species  there  are  two  nearly  equal,  long, 
slender  adambulacral  spines  to  each  plate  proximally  as  well  as 
distally. 

Another  arctic  species  (U.  pcmopla  Stuxberg)1  from  Spitzbergen, 
etc.,  has  more  resemblance,  for  it  has  few  dorsal  spines  and  abundant 
dorsal  groups  of  minor  pedicellariae.  But  these  pedicellariae  are  not 
of  the  same  form  and  are  much  smaller,  as  also  the  major  pedicel- 
lariae. Its  ventral  spines  are  much  more  numerous,  longer  and  more 
slender. 

ORTHASTERIAS  LEPTOLENA  Verrill,  sp.  nov. 

Plate  LXIV,  figures  i,  la,  2,  20   (ventral  and  dorsal  sides  of  types)  ;  plate 
LXXVII,  figures  2,  a-d  (pedicellariae). 

The  disk  is  small;  the  rays  five,  long,  slender,  tapered,  well 
rounded.  The  radii  of  the  larger  (No.  i8a)  are  7  mm.  and  76  mm. ; 
ratio,  nearly  i :  n.  The  smaller  has  radii  of  4  mm.  and  44  mm.; 
ratio,  i:  ii. 

The  dorsal  spines  of  the  rays  and  disk  are  rather  long  and  of 
nearly  uniform  size  and  length,  numerous  but  not  crowded,  mostly 
one  to  a  plate,  and  pretty  uniformly  distributed,  nearly  in  quincunx, 
or  in  five  indistinct,  alternating  rows  on  the  rays,  the  median  row  not 
differing  from  the  rest.  These  spines  are  tapered,  but  obtuse  and 
rough  or  subsulcate  at  the  tips,  which  are  often  flattened.  Between 
them  there  are  large,  denticulate  dermal  pedicellarise,  often  about 
half  as  thick  as  the  spines.  Papular  areas  are  large;  the  papulae 
small  and  numerous. 

The  superomarginal  spines  form  a  single  regular  row,  one  to  all 
plates.  They  are  similar  to  the  dorsals  in  size  and  form.  The  infero- 
marginals  form  a  very  distinct  double  row,  two  to  a  plate,  sepa- 
rated from  the  upper  ones  by  a  rather  deep  naked  channel,  with 
large  papular  pores.  The  inferomarginal  spines  are  like  the  upper 
ones  in  form,  but  a  little  longer  and  less  tapered,  obtuse.  There  are 
no  spiniferous  interactinal  plates.  The  peractinal  row  is  represented 
by  a  proximal  row  of  small,  narrow,  spineless  plates,  five  to  ten  in 
number,  on  the  different  rays,  and  joined  closely  to  the  adambulacrals. 
In  the  smaller  specimen  these  actinal  ossicles  are  rudimentary  or 
lacking. 

1  See  Doderlein,  1900,  p.  204,  pi.  iv,  fig.  2 ;  pi.  v,  figs.  I,  2. 


SHALLOW-WATER   STARFISHES  183 

Adambulacral  spines  are  irregularly  diplacanthid,  small,  slender, 
terete  or  obtuse,  about  half  as  long  as  the  inferomarginals  and 
scarcely  one-third  as  thick.  The  outer  ones  are  a  little  longer  and 
often  slightly  clavate ;  the  proximal  ones  become  distinctly  longer  and 
more  tapered.  There  is  an  unusually  wide  naked  space  between  the 
adambulacral  and  the  inferomarginal  spines. 

Oral  and  epioral  spines  longer  and  distinctly  stouter. 

Remarkably  large  dermal  major  pedicellariae  are  sparingly  scat- 
tered over  the  back  and  sides.  They  are  mostly  of  two  forms ;  the 
larger  are  stout,  erect,  compressed,  ovate,  with  a  stout  base,  often 
nearly  as  thick  as  the  adjacent  spines ;  apex  a  little  acuminate,  obtuse. 
The  others  are  about  as  long,  but  not  so  stout,  compressed,  con- 
tracted in  the  middle,  with  the  blades  a  little  spatulate  and  dentate 
or  unguiculate  at  the  tips.  Much  smaller,  simple,  lanceolate  ones 
also  occur.  Thick  wreaths  of  rather  large  minor  pedicellariae  sur- 
round the  basal  half  of  the  dorsal  and  superomarginal  spines ;  on  the 
inferomarginal  spines  they  form  secund  groups. 

The  madreporite  is  rather  prominent,  with  very  fine  gyri ;  it  is  not 
surrounded  by  special  spines. 

The  color,  as  dried,  is  dark  purplish  or  greenish  brown  above ;  pale 
yellowish  below ;  madreporite  pale  lemon-yellow. 

The  smaller  specimen  agrees  well  with  the  larger,  except  in  having 
all  the  spines  disproportionately  smaller,  and  in  lacking  visible  per- 
actinal  plates  and  large  dorsal  dermal  pedicellariae;  but  similar  ones 
occur  beneath,  on  the  interradial  areas.  Minor  pedicellariae  form 
large  wreaths  on  the  dorsal  spines. 

Two  specimens  (No.  18)  were  dredged  in  shallow  water  in 
Departure  Bay,  British  Columbia,  by  Prof.  John  Macoun  and  party, 
of  the  Canadian  Geological  Survey,  in  1909. 

These  specimens  are  probably  both  young,  though  the  larger  is 
nearly  as  large  as  the  type  of  O.  kcehleri,  with  which  it  appears 
nearly  to  agree  in  proportions.  It  may,  possibly,  be  the  young  of 
O.  columbiana,  which  comes  from  the  same  district ;  but  the  smallest 
undoubted  specimens  of  that  species  that  I  have  seen  have  much 
stouter  rays  and  much  larger  and  less  numerous  spines,  though  the 
rays  are  scarcely  longer,  and  it  has  a  row  of  spiniferous  peractinal 
plates.  The  large  dermal  dorsal  pedicellariae  are  similar  in  the  two 
forms,  but  relatively  smaller  and  less  stout  in  the  young  columbicma. 
The  minor  pedicellariae  are  also  smaller  in  the  latter  and  form 
smaller  wreaths  on  the  dorsal  spines.  Therefore  it  is  more  probable 
that  they  are  distinct,  though  closely  related  species.  It  would 
certainly  be  unwise  to  unite  them  without  intermediate  forms. 


184  VERRILL 

ORTHASTERIAS  GONOLENA  Verrill,  sp.  nov. 

Plate  LXVII  (dorsal  side)  ;  plate  LXVIII,  figure  i   (actinal  side)  ;  plate  LXIX, 

figure  2  (portion  enlarged)  ;  plate  LXXXII,  figures  3-36,  4-40  (details). 

Rays  five,  of  moderate  length,  rather  stout,  somewhat  angular. 
Disk  small.  Radii  of  the  specimen  figured,  8  mm.  and  68  mm.; 
ratio,  i :  8.5.  Radii  of  a  larger  specimen,  13  mm.  and  83  mm. ; 
ratio,  i :  6.4. 

Dorsal  spines  rather  long,  tapered,  obtuse,  not  much  grooved  or 
fluted,  well  spaced.  They  form  three  rows,  quite  regular  in  the 
smaller  specimens,  less  so  in  the  larger.  The  median  row  is  distinct, 
having  more  regular  and  more  numerous  spines,  though  they  are  not 
longer.  The  spines  stand  singly  on  rather  stout,  alternate  plates. 
They  are  surrounded  in  the  dry  specimens  by  a  large  subbasal  wreath 
of  small  minor  pedicellariae  attached  to  the  edge  of  a  thick  basal 
sheath.  Papular  areas  unequal;  some  are  very  small,  others  large, 
with  many  small  papulae.  Superomarginal  spines  similar  to  the  dor- 
sals in  size  and  form.  They  form  a  regular  row,  one  to  each  alternate 
plate.  No  intermarginal  spines;  interactinal  plates  are  sporadic, 
spineless,  small,  compressed,  standing  edgewise,  scarcely  visible 
unless  cleaned  by  potash.  The  integument  is  rather  thick. 

The  inferomarginal  spines  stand  two  to  a  plate,  placed  obliquely 
in  two  rows,  close  to  the  adambulacrals,  but  with  a  row  of  small, 
single  papulae  between.  These  spines  are  about  a,s  long  as  the  supero- 
marginals,  but  not  so  stout,  and  they  are  decidedly  flattened  dis- 
tally.  There  is  a  wide,  conspicuous  intermarginal  channel,  in  which 
papular  areas  of  small  size  alternate  with  the  plates  in  a  regular  row. 
In  this  lane  or  channel  are  also  scattered  erect,  stout,  wedge-shaped 
pedicellariae  of  large  size,  often  nearly  as  stout  as  the  spines.  The 
lane  is  crossed  by  the  long,  stout,  descending  apophyses  of  the  su- 
peromarginal  plates,  which  meet  and  overlap  the  ascending  lobes  of 
the  inferomarginals,  without  an  intermediate  ossicle.  The  alternate 
spineless  plates  have  equally  large  apophyses.  The  dorsal  connec- 
tive ossicles  are  stout,  nearly  as  wide  as  the  plates. 

The  adambulacral  spines  stand  obliquely,  two  to  a  plate,  erect  in 
the  type,  nearly  equal  in  length,  the  inner  ones  more  slender,  all 
distinctly  flattened  and  sometimes  slightly  grooved  distally,  except 
the  adoral  ones,  which  are  longer,  more  slender,  terete  or  nearly  so. 

The  dorsal  dermal  pedicellariae  are  few  on  the  type,  wedge-shaped. 
Some  very  large  ones  are  found  on  the  sides,  as  mentioned  above, 
and  some  of  the  same  form  and  equally  large  are  sometimes  found 
attached  to  the  inner  edge  of  the  adambulacral  groove,  or  to  the  bases 


SHALLOW-  WATER   STARFISHES  185 

of  the  spines  ;  these  are  usually  denticulate.     Smaller  short-ovate 
major  pedicellarige  occur  on  the  interradial  areas  (see  figures). 

Specimen  described  and  figured  is  from  off  southern  California 
(Professor  Ritter,  coll.  Yale  Museum). 

I  have  also  studied  specimens  of  this  species  in  the  Museum  of 
Comparative  Zoology  from  the  Gulf  of  California  (No.  1214),  eight 
young,  from  35  mm.  to  75  mm.  in  diameter;  La  Paz  (No.  1215),    r 
one  small;  Santa  Cruz  Island,  California  (No.  1212),  one;  Santa  "1  ^ 
Barbara,  California  (No.  1413),  about  150  mm.  in  diameter;  San 


Diego,  California  (No.  1825),  one.     TH^tX-oV  C+4yn 

This  species  differs  from  O.  caHfornica  and  O.  columbiana,  both 
of  which  it  resembles,  in  lacking  peractinal  spines.  Its  dorsal  spines 
are  not  fluted  like  those  of  columbiana,  nor  do  they  have  the  stout, 
elevated  sheath  carrying  the  pedicellariae,  present  in  that  species; 
the  large  wreaths  in  this  are  basal.  The  pedicellariae  are  also  dis- 
tinctive. The  rays  are  more  angular  and  the  ossicles  stouter. 

Genus  Distolasterias  Perrier. 

Distolasterias  (pars)  PERKIER,  op.  cit,  1896,  p.  34.    Type,  D.  stichantha  Sladen, 
from  Japan.    See  above,  p.  47. 

This  genus,  as  restricted  to  the  stichantha  group,  is  closely  related 
to  Orthasterias.  It  differs  mainly  in  having  no  interactinal  plates, 
visible  externally,  at  least  in  our  species,  even  after  removal  of  the 
dermis  by  potash,  though  a  rudiment  may,  perhaps,  exist  internally. 
Also  in  having  more  numerous  rows  of  dorso-lateral  plates  and 
spines,  and  in  having  the  plates  more  closely  joined,  mostly  by  their 
own  overlapping  lobes,  with  only  few  small  connective  ossicles. 
Thus  the  papular  areas  are  small  and  numerous,  with  papulae  in 
small  groups.  The  superomarginal  and  intermarginal  lanes  are 
narrow,  each  with  a  regular  row  of  small  papular  areas,  with  few 
papulae.  The  inferomarginal  lane  is  very  narrow,  with  the  papulae 
standing  singly  or  in  pairs.  Alternate  superomarginals  are  not 
spineless  in  our  species. 

DISTOLASTERIAS  CHELIFERA  Verrill,  sp.  nov. 
Plate  LXXXI,  figures  i-ifc  (spines  and  pedicellariae)  ;  plate  ex,  figures  I,  2  (type). 

A  good  dry  specimen  of  this  species  (No.  1346)   has  the  radii     *CV>-  */ 
10  mm.  and  100  mm.  ;  ratio,  i  :  10.    A  larger  one,  No.  19,  from  British  1  ""' 
Columbia,  has  the  radii  16  mm.  and  180  mm.  ;  ratio,  1:11.2. 

The  five  rays  are  long,  slender,  nearly  terete,  gradually  tapered 
to  unusually  slender  tips,  with  a  small  apical  plate. 


1 86  VERRILL 

Dorsal  spines  are  numerous,  pretty  evenly  distributed,  and  mostly 
stand  singly,  arranged  apparently  in  quincunx ;  or  they  might  be  said 
to  form  nine  alternating  rows,  with  some  intermediate.  Dorsal 
median  row  distinct,  but  scarcely  different  in  size.  The  dorsal  spines 
are  rather  short,  for  a  Distolasteiias,  and  often  a  little  clavate  and 
deeply  fluted  near  the  obtuse  tips.  Papular  areas  are  mostly  rather 
large  with  few  papulae.  Minor  pedicellariae  are  very  small  and  form 
dense  wreaths  around  the  middle  of  the  spines. 

Superomarginal  spines  are  like  the  dorsals,  but  a  little  longer; 
they  form  a  regular  row,  one  to  a  plate.  Inferomarginals  rather 
more  slender,  two  to  a  plate,  mostly  flattened  and  slightly  gouge- 
shaped  at  the  truncate  tips,  and  with  large  clusters  of  minor  pedicel- 
lariae. Adambulacrals  two  to  a  plate,  nearly  equal,  in  two  equal, 
regular,  divergent  rows,  slender,  not  flattened.  The  proximals  and 
epiorals  longer;  apical  oral  pair  of  spines  stouter.  The  peractinal 
plates  are  small  and  spineless  or  lacking. 

In  the  lane  between  the  adambulacral  and  inferomarginal  spines 
there  is  a  regular  row  of  small  papular  areas,  each  having  usually 
a  single  papula.  A  similar  series  exists  between  the  inferomarginals 
and  superomarginals. 

Dorsal  major  pedicellariae  are  mostly  large,  elongated,  sometimes 
nearly  as  stout  as  the  spines,  usually  decumbent,  with  spatulate  jaws 
strongly  unguiculate  with  one  to  three  curved  denticles  at  the  tip. 
Some  are  stouter,  erect,  more  or  less  ovate.  The  denticles  of  the 
dorsal  major  pedicellariae  are  few,  large  curved  and  interlocking, 
sometimes  only  one  fitting  between  two  on  the  opposite  valve ;  often 
two  on  each;  or  two  on  one  and  three  on  the  other.  Similar  large 
ones  occur  on  the  sides  and  on  the  actinal  interradial  areas ;  those  of 
the  latter  areas  are  often  more  elongated  and  remarkably  large. 

The  adambulacral  major  pedicelleriae  are  more  slender,  lanceolate, 
acute. 

The  type  is  from  Vancouver  Island  (No.  1346,  Mus.  Comp.  Zool.). 
The  larger  specimen  (No.  19,  Geological  Survey  of  Canada)  is  from 
British  Columbia. 

This  species  is  notable  for  its  unusually  numerous  fluted  dorsal 
spines  and  very  large  unguiculate  pedicellariae. 

The  larger  dry  specimen  mentioned  above  agrees  very  well  with 
the  type,  except  in  features  due  to  its  greater  age.  The  dorso-lateral 
spines  are  more  numerous  and  cannot  be  said  to  stand  in  any  definite 
rows;  their  arrangement  is  in  irregular  quincunx,  or  in  irregular 
oblique  transverse  series  of  about  five  on  each  side  of  the  slightly 


SHALLOW- WATER   STARFISHES  187 

more  prominent  carinals.  The  dorso-lateral  plates  are  openly  stellate- 
reticulate,  with  rather  narrow  connective  ossicles,  leaving  large  angu- 
lar papular  areas  with  many  papulae.  Each  plate  bears  one  spine 
on  a  raised  central  boss. 

Minor  pedicellariae  are  very  abundant  between  and  around  the 
bases  of  the  spines.  Marginal  and  adambulacral  spines  are  nearly 
as  in  the  type.  No  peractinal  plates  are  visible.  The  specimen  is 
dry  and  varnished.  It  was  stained  dark  purple  to  imitate  the  natural 
color. 

British  Columbia  (Geological  Survey  of  Canada,  No.  19). 

Genus  Urasterias  Verrill. 
Plate  LXX,  figures  1-4. 

Urasterias  VEWULL,  Amer.  Journ.  Sci.,  xxvni,  p.  67,  1909.    Type,  U.  line  kit 
(M.  and  T.). 

Large  starfishes  with  a  loose,  openly  reticulated  dorsal  skeleton, 
composed  largely  of  slender  ossicles,  with  few  larger  plates,  mainly 
in  the  interradial  or  genital  areas.  Dorsal  spines  large,  mostly  iso- 
lated on  the  plates;  median  radial  row  usually  distinct;  elsewhere 
they  are  mostly  scattered.  Superomarginal  spines  elongated,  mostly 
standing  in  single  rows.  No  notable  actinal  plates  nor  spines.  Adam- 
bulacral plates  diplacanthid  or  subdiplacanthid. 

Some  details  of  the  type  are  figured  on  pi.  LXX,  figs.  1-4. 

Genus  Parastcrias  Verrill,  nov.    See  also  p.  53. 

Type,  P.  albertensis  Ver. 

Dorsal  and  marginal  skeleton  fairly  well  developed,  reticulated 
nearly  as  in  typical  Asterias.  Interactinal  plates  lacking  or  rudi- 
mentary, not  showing  externally.  Adambulacral  plates  diplacanthid. 

PARASTERIAS  ALBERTENSIS  Verrill,  sp.  nov. 
Plate  LVII,  figures  i,  2;  plate  LXX,  figure  6. 

A  five-rayed  species  with  rather  depressed  rays  and  openly  reticu- 
late structure,  the  dorsal  ossicles  being  thin  and  weak,  while  the 
papular  areas  are  large  and  conspicuous.  Dorsal  spines  are  not 
numerous,  very  small,  tapered,  obtuse,  not  capitate,  pretty  evenly 
distributed  on  the  ossicles,  standing  singly  and  not  forming  a  con- 
spicuous median  row. 

Madreporite  rather  large,  convex,  fine-grained,  not  encircled  by 
spines.  Minute  major  pedicellariae,  acute-ovate  in  form,  are 


1 88  VERRILL 

sparsely  scattered  on  the  papular  areas.  The  distal  dorsal  spines 
have  thick  wreaths  of  minor  pedicellariae. 

There  are  but  two  distinct  double  rows  of  lateral  and  ventral 
spines.  The  upper  one,  which  appears  to  be  the  superomarginal, 
usually  has  three  spines  to  a  plate  proximally  but  only  two  distally ; 
the  spines  are  like  the  dorsals  but  slightly  larger.  The  lower  or 
inferomarginal  row  bears  two  spines  to  a  plate,  regularly,  the  spines 
being  like  those  above.  This  row  of  plates  is  separated  from  the 
adambulacrals  by  a  row  of  papular  areas. 

The  adambulacral  plates  have  each  two  slender,  divergent  spines, 
which  are  tapered  and  subacute  and  carry  numerous  acute-lanceo- 
late major  pedicellariae  of  moderate  size;  similar  pedicellariae  are 
abundant  on  the  interradial  areas  and  marginal  channels. 

The  radii  are  13  mm.  and  54  mm. ;  ratio,  i :  4. 

Albert  Bay,  British  Columbia  (Miss  Kate  Foote,  January,  1887, 
Yale  Museum).  &  ttty 

This  species  looks  more  like  the  typical  species  of  Asterias  of  the 
North  Atlantic  than  any  other  west-coast  species  known  to  me.  It 
resembles  some  varieties  of  A.  rubens  and  A.  forbesi  quite  strongly, 
but  the  evenly  reticulated  arrangement  of  the  dorsal  plates  and 
spines  and  the  absence  of  a  larger  median  row  give  it  a  character- 
istic appearance.  The  lack  of  interactinal  plates  and  spines  is  dis- 
tinctive. 

Genus  Allasterias  Verrill. 

Type,  A.  rathbuni  Verrill. 
Allasterias  VERRILL,  Atner.  Journ.  Sci.,  xxvm,  p.  65,  1909. 

Distinguished  by  the  arrangement  of  the  adambulacral  spines,  in 
several  series,  of  which  one  is  deeper  within  the  groove  on  alternate 
plates.  Disk  rather  large,  areolate.  Dorsal  ossicles  numerous,  but 
small,  arranged,  both  on  the  disk  and  rays,  in  a  reticulate  manner 
around  the  papular  areas,  which  are  numerous,  and  bear  large  groups 
of  small  papulae.  Spines  numerous,  arranged  irregularly,  or  placed 
around  the  papular  areas,  but  usually  forming  a  median  radial  series. 
Upper  marginal  plates  rather  large  and  stout,  so  as  to  form  an 
angular  margin,  bearing  one  or  several  spines  larger  than  the  dorsals. 
Lower  marginals  not  close  to  the  adambulacrals,  bearing  in  the  type 
two  or  three  spines,  longer  than  the  upper  ones.  Actinals  rudi- 
mentary or  lacking.  A  wide  intermarginal  channel,  with  many  for- 
ficulate  pedicellariae. 


SHALLOW-WATER  STARFISHES  189 

The  arrangement  of  the  adambulacral  spines  is  peculiar,  for  while 
the  plates  bear  alternately  two  and  three  spines,  the  inner  spine  of 
the  alternate  plates  is  attached  deep  within  the  groove  on  a  special 
lobe  of  the  inner  edge  of  the  plate,  while  the  two  outer  ones  stand 
on  the  actinal  end,  nearly  in  line  with  the  spines  of  the  adjacent 
actinal  plates ;  thus  they  seem  to  form  four  or  five  rows.  Jaw-plates 
bear  two  pairs  of  apical  oral  spines,  one  above  the  other. 

The  interradial  areas,  near  the  jaws,  are  without  spines  and  in  the 
dry  specimens  they  are  sunken  in  the  form  of  pits.  The  position  of 
the  genital  pores  was  not  determined  for  lack  of  alcoholic  specimens. 

ALLASTERIAS  RATHBUNI  Verrill. 

Plate  LXXVIII,  figure  2  (details),  var.  nortonensis. 

Allasterios  rathbuni  VERRILL   (pars),  Amer.  Journ.  Sci.,  xxvra,  p.  65,  1909, 

figs.  5,  6,  7  (as  varieties). 
Asterias  rubens  ?  MURDOCH,  Rep.  Int.  Polar  Expedition  to  Point  Barrow,  p. 

159,   1885    (MOM  Linne). 

Rays  five,  broad  at  base  and  rapidly  tapering  to  acute  tips.  Radii, 
25  mm.  and  100  mm. ;  ratio,  1 : 4.  Small  major  pedicellariae  are 
abundant  all  over  the  dorsal  and  lateral  surfaces. 

Dorsal  skeleton  rather  feeble,  composed  of  a  great  number  of 
small  ossicles,  arranged  in  an  openly  reticulated  manner,  so  that  the 
texture  is  rather  soft  and  flaccid,  in  alcohol.  The  dorsal  papular 
areas  are  numerous  and  contain  many  small  papulae.  Disk  rather 
broad,  but  probably  abnormally  so  in  the  dried  specimens,  owing  to 
the  flattening  when  soft. 

The  whole  dorsal  surface  is  conspicuously  areolate  or  reticulate, 
the  areolations  mostly  1.5  mm.  to  2  mm.  broad.  The  dorsal  spines 
are  very  small  and  numerous,  sometimes"  almost  like  round  or  capi- 
tate granules,  being  scarcely  higher  than  thick,  but  in  other  examples 
clavate  or  partly  acute ;  they  are  scattered  or  arranged  in  single  rows 
on  all  the  ossicles,  so  as  to  form  a  border  around  the  papular  areas ; 
toward  the  sides  of  the  rays  they  are  distinctly  longer  and  mostly 
clavate  or  subacute. 

The  superomarginal  spines  form  a  wide  band  of  small,  crowded 
spines,  five  to  ten  or  more  on  a  plate.  They  are  mostly  larger  and 
longer  than  the  dorsals,  and  two  or  three  times  as  long  as  thick, 
mostly  cylindrical  or  clavate,  sometimes  gouge-shaped.  Below  this 
band  there  is  a  broad  intermarginal  channel  with  large  papular  areas 
and  numerous  rather  large,  pointed  major  pedicellariae.  This  chan- 
nel rapidly  widens  at  the  bases  of  the  rays. 


IQO  VERRILL 

The  lower  marginals  form  a  double  row,  mostly  two  to  a  plate; 
they  are  similar  to  the  upper  ones,  but  longer  and  mostly  more 
clavate,  often  with  slightly  gouge-shaped  tips.  Between  the  upper 
and  lower  marginals,  at  the  bases  of  the  rays,  a  short  intermdiate 
row  of  ossicles  is  sometimes  interpolated,  and  defined  by  a  row  of 
small  papular  pores ;  their  spines  are  like  the  lower  marginals,  and 
mostly  stand  singly.  Synactinal  spines  are  lacking ;  there  is  a  wide, 
naked  channel  between  the  adambulacrals  and  inferomarginals. 

The  adambulacral  spines  are  peculiarly  arranged,  and  look  as  if 
in  four  rows,  in  adult  specimens.  Each  plate  bears  two  spines,  or 
alternately  two  and  three ;  but  on  the  alternate  plates,  having  three 
spines,  the  inner  spine  is  set  on  the  inner  edge  of  the  plate,  deep 
within  the  groove,  while  its  outer  spines  are  at  the  summit,  nearly 
but  not  quite  in  line  with  the  spines  of  the  alternate  plates.  The 
outer  of  these  spines  are  situated  further  back,  at  the  outer  end  of  the 
plates.  These  spines  are  not  very  slender,  either  tapered  or  clavate, 
mostly  obtuse,  as  long  as  the  marginals  or  longer,  and  more  slender. 
Many  of  the  inner  ones  bear  small,  acute  major  pedicellarise ;  the 
outer  ones  have  a  small  cluster  of  minor  pedicellariae. 

The  apical  jaw-plate  bears  an  inner  close  pair  of  stout,  tapered 
spines,  low  down  within  the  mouth,  and  a  similar  upper  pair  at  the 
upper  margin;  both  pairs  bear  several  large  major  pedicellariae. 

Major  or  forficulate  pedicellariae  are  usually  everywhere  abundant, 
scattered  over  the  surface,  between  the  dorsal,  marginal,  and  actinal 
spines,  and  especially  on  the  lateral  channels  and  interradial  areas. 
The  larger  ones  are  compressed,  rather  large,  lanceolate  or  acute- 
triangular,  with  a  sharp  or  acuminate  apex.  Those  that  are  scat- 
tered on  the  dorsal  surface  are  smaller,  unequal  in  size,  but  similar 
in  form,  though  less  acute.  The  larger  and  longer  ones  measure 
mostly  about  i.oo  mm.  X  0.33  mm.  to  0.37  mm. ;  the  stouter  ones, 
0.87  mm.  X  0.37  mm.  and  0.72  mm.  X  0.39  mm. 

Minor  pedicellariae,  of  very  small  size,  are  attached  to  many  of  the 
spines  in  small  groups,  and  are  also  found  scattered  on  the  dorsal 
papular  areas. 

The  madreporic  plate  is  large,  convex,  with  numerous  gyri. 

Dedicated  to  Dr.  Richard  Rathbun  of  the  U.  S.  National  Museum. 

The  type  specimens  are  from  Maloska  (Professor  Kincaid). 
'  This  species  is  so  different  from  any  other  known  to  me  from  this 
region  that  no  detailed  comparisons  seem  necessary.     Its   finely 
and  regularly  areolated  surface,  flaccid,  reticulated  dorsal  skeleton, 
and  small  spines  will  serve  to  distinguish  it  at  a  glance. 


SHALLOW-WATER   STARFISHES  19! 

The  arrangement  of  the  adambulacral  spines  is  so  peculiar  that  it 
should,  even  on  that  account  alone,  form  the  type  of  a  genus.  But  it 
is  also  peculiar  in  the  structure  of  the  dorsal  skeleton,  the  character 
of  the  marginal  spines,  etc. 

The  only  other  species  known  to  me,  besides  the  following  four, 
described  as  having  a  similar  arrangement  of  a  row  of  spines  within 
the  groove,  is  A.  versicolor  (Sladen),  from  off  Kobe,  Japan.  In 
that  species  the  alternate  plates  bear  only  one  spine,  and  the  dorsal 
and  lateral  spines  are  larger,  much  less  numerous,  and  differently 
arranged.  It  evidently  belongs  to  the  same  genus. 

Besides  A.  versicolor,  there  is  another  species,  A.  amurensis  Liit- 
ken,  as  Asterias,  from  further  north,  on  the  Asiatic  coast,  and  also 
recorded  from  Yokohama  by  Sladen,  that  must  bear  considerable 
resemblance  to  our  species,  though  the  dorsal  and  upper  marginal 
spines  are  fewer  and  longer.  Liitken,  however,  does  not  mention  any 
such  peculiar  arrangement  of  the  adambulacral  spines  as  is  found  in 
this  group.  I  have  had  no  opportunity  .to  study  his  species. 

A  small  Korean  starfish,  described  by  Sladen  (1878,  p.  432),  under 
the  name  of  Asterias  rubens,  var.  migratum,  probably  belongs  to  this 
genus,  and  should  be  called  Allasterias  migrata,  but  its  immaturity 
(greater  radius  of  the  larger  specimen,  16  mm.)  renders  it  impossible 
to  determine  whether  it  be  distinct  from  the  other  known  species, 
until  a  series  can  be  studied. 

ALLASTERIAS  RATHBUNI   NORTONENSIS  Verrill. 

Plate  LXXVIII,  figure  2  (details  of  type)  ;  text-figures  Nos.  8,  9. 
Allasterias  rathbuni,  var.  nortonensis  VERBILL,  op.  cit.,  1909,  p.  66,  fig.  7. 

Rays  five,  broad  at  base,  depressed,  acute.  Radii  of  the  type  are 
22  mm.  and  82  mm. ;  ratio,  1 : 3.8. 

The  dorsal  spines  are  numerous,  pretty  evenly  spaced  and  scat- 
tered over  the  rays  and  disk  on  the  openly  reticulated,  slender 
ossicles.  They  are  partly  blunt,  cylindric,  and  partly  clavate,  but  not 
strongly  so,  rather  longer  than  in  the  typical  form.  Papular  areas 
are  large,  with  many  small  papulae  and  scattered  small  dermal  pedi- 
cellariae.  Madreporic  plate  large  and  strongly  convex.  The  dorsal 
spines  bear  but  few  minor  pedicellariae.  Some  larger  acute  major 
pedicellariae  occur  among  the  spines. 

The  adambulacral  spines  stand  mostly  two  and  three  to  a  plate  in 
alternation.  The  inner  or  furrow  spines  are  elongated,  tapered,  sub- 
acute  and  often  bear  a  cluster  of  small  major  pedicellariae.  The  outer 


192 


VERRILL 


ones  are  a  little  stouter  and  less  acute,  mostly  not  quite  so  long. 
They  usually  bear  a  small  group  of  minor  pedicellariae  on  the  outer 
side. 

The  inferomarginal  spines  mostly  stand  two  or  three  to  a  plate, 
forming  a  pretty  regular  double  row.  They  are  much  stouter  than 
the  adambulacrals  and  not  quite  so  long ;  many  are  a  little  flattened 
or  pinched  up  at  the  tip,  and  often  slightly  gouge-shaped. 

The  superomarginals  are  more  numerous  and  shorter,  of  similar 
shapes.  Near  the  disk  they  often  stand  four  to  six  on  a  plate, 
crowdedly.  There  are  no  peractinal  plates.  Major  pedicellariae  are 
numerous  on  the  sides  and  actinal  surface.  The  larger  of  these  are 


FIG.  8. 


Allastcrias  rathbiini  nortonensis  Ver.     No.  7621,  U.  S.  Nat.  Mus.    Spines;  a,  intra-ambu- 
lacral;  b,  outer-adambulacral ;  c,  inferomarginal.      X  20. 

elongated,  acute  or  acuminate  in  profile,  with  the  valves  mostly 
narrow  spatulate ;  some  of  them  are  dentate  at  the  tips,  with  two  or 
three  curved  teeth.  The  larger  ones  measure  0.67  mm.  X  0.30  mm. ; 
0.67  mm.  X  0.27  mm. ;  0.65  mm.  X  0.25  mm. ;  0.57  mm.  X  0.23  mm. 
Many  are  smaller,  short  and  thick,  sometimes  as  broad  as  long. 
These  measure  0.50  mm.  X  0.30  mm. ;  0.40  mm.  X  0.27  mm. ; 
0.35  mm.  X  0.40  mm. ;  and  smaller. 

The  type,  and  the  only  specimen  seen,  was  from  Norton  Sound 
(coll.  Murdoch,  No.  7621,  U.  S.  Nat.  Mus.).  This  presents  several 
rather  strongly  marked  characters,  which  might  be  considered 
specific  if  persistent  in  a  large  series  of  specimens.  But  with  the 
small  number  of  specimens  available  for  study  it  seems  better  to 
consider  it  a  subspecies  at  present.  Very  likely  the  species  may  prove 
to  be  a  very  variable  one.  The  characters  that  seem  most  important 
are  the  smaller  size  and  stouter  form  of  the  major  pedicellariae,  and 


SHALLOW-WATER   STARFISHES 


193 


the  different  forms  of  the  minor  pedicellariae.  The  dorsal  and  mar- 
ginal spines  are  longer  than  in  the  typical  form,  and  the  papulae  are 
more  numerous. 


FIG.  9. 

Allajterias  rothbvni  nortonensis  Ver.  Pedicellarue  from  the  axillary  or  actinal  interradial 
areas;  a-c,  acute  lanceolate  forms;  d,  e,  m,  the  same  with  the  valves  spatulate  and  dentate; 
g-l,  small  short  forms.  X  32. 

ALLASTERIAS  ANOMALA  Verrill. 

Plate  LIX,  figure  2  (type,  actinal  side)  ;  plate  LX,  figure  2  (type,  dorsal)  ;  plate 
LXIX,  figure  5  (type,  actinal  X  5%)  ;  plate  LXXVII,  figure  3;  plate  LXXVIII, 
figures  3,  4  (pedicellariae  and  spines).  ~) 

dllasterias  rathbuni,  var.  anomala  VERRILL,  op.  cit,  1909,  p.  66,  figs.  5,  6. 

Rays  five,  broad  at  base,  regularly  tapered  to  rather  acute  tips,  and 
margined  on  the  sides  with  a  fringe  of  superomarginal  spines.  Radii 
of  the  type,  23  mm.  and  87  mm. ;  ratio,  i :  3.8.  Breadth  of  rays  at 
base,  29  mm. 

The  dorsal  ossicles  are  slender  and  loosely  reticulated,  leaving 
large  papular  areas,  with  numerous  papulae.  The  dorsal  spines  are 
short,  small  and  numerous,  not  crowded,  clavate  and  capitate.  They 
mostly  stand  isolated  on  the  ossicles  around  the  papular  areas,  often 
forming  imperfect  circles  or  polygons  having  twelve  to  twenty 
spines  in  the  groups,  so  that  they  present  a  somewhat  reticulated 
arrangement.  A  median  radial  row  is  distinct,  due  to  the  increased 
number  of  spines,  not  to  their  larger  size. 

The  madreporic  plate  is  large,  round,  and  convex,  without  any 
special  spines  around  it. 

The  superomarginal  spines  form  a  conspicuous  marginal  fringe, 
owing  to  their  numbers  rather  than  to  their  size.  They  are  short, 
14 


194  VERBILL 

..  .  J  stout,  enlarged  distally,  truncate,  and  mostly  grooved  or  gouge- 
shaped  at  the  end.  They  are  much  crowded  and  stand  mostly  five  or 
six  to  a  plate. 

The  inferomarginals  are  similar  in  form,  a  little  longer,  and  mostly 
stand  two  or  three  to  a  plate,  in  oblique  groups.  There  is  a  well 
marked  channel  between  these  and  the  superomarginals,  but  no 
intermarginal  rows  of  spines.  A  similar  regular  channel  separates 
the  inferomarginals  from  the  adambulacrals.  These  channels  are 
both  occupied  by  large  papular  areas,  with  clusters  of  small  papulae, 
and  by  rather  large,  acute  major  pedicellariae. 

The  adambulacral  spines  stand  mostly  two  and  three  to  a  plate, 
alternately,  every  other  plate  bearing  a  rather  long,  tapered,  furrow- 
spine.  Most  of  their  other  spines,  on  the  actinal  side,  are  stout, 
enlarged  distally,  and  grooved  or  gouge-shaped  on  the  outer  side. 

The  minor  pedicellariae  are  few  and  very  small.  They  are  entirely 
lacking  on  most  of  the  dorsal  spines,  and  very  few  occur  on  the 
marginal  spines. 

The  major  pedicellariae  are  frequent  both  on  the  actinal  and  on  the 
lateral  areas,  but  not  abundant.  They  are  few  and  scattered  on  the 
dorsal  surface.  The  larger  are  compressed,  acute-lanceolate,  some 
more  acute  than  others;  the  smaller  ones  are  short-lanceolate  or 
ovate. 

The  jaws  are  elongated  and  rather  compressed.  The  two  upper 
apical  spines  are  stout  and  bear  small  clusters  of  acute  major  pedi- 
cellariae; the  lower  ones  are  much  smaller,  bent,  acute.  There  are 
two  spaced  pairs  of  stout  epiorals. 

The  type  is  from  St.  Michael,  Alaska  (L.  M.  Turner,  1874-76, 
No.  3821,  U.  S.  Nat.  Mus.).  Vtt-vifw 

ALLASTERIAS  FORFICULOSA  Verrill,  sp.  nov. 

;*     • 

Plate  LXXXIII,  figures  3-3*:  (details)  ;  plate  LXXXIV,  figure  i,  a-h   (pedicel- 
lariae) ;  text-figures  10,  n. 

ft 

Asterias  amurensis  IVES,  Proc.  Acad.  Nat  Sci.,  Philad.,  for  1891,  p.  212,  plate 
vm,  figures  5-8  (non  Liitken). 

Rays  five,  rather  short,  wide  at  base,  depressed,  tapering  rapidly, 
rather  acute,  bordered  by  a  fringe  of  superomarginal  spines.  Radii, 

20  mm.  and  58  mm. ;  ratio,  about  1:3;  breadth  of  rays  at  base, 

21  mm. 

The  dorsal  spines  are  well  spaced,  rather  scattered,  but  on  the 
sides  of  the  rays  forming  pretty  evident  radial  rows;  median  row 
distinct  but  not  prominent.  They  are  short,  stout,  about  twice  as 


SHALLOW-WATER  STARFISHES 


195 


high  as  thick,  mostly  cylindric  or  slightly  clavate,  obtuse,  but  some 
are  acute.  They  are  mostly  surrounded  by  small  wreaths  of  very 
small  minor  pedicellariae. 

The  dermal  surface  is  notably  smooth  and  even.  The  papular 
pores  are  very  small,  not  very  numerous.  Very  acute  lanceolate 
major  pedicellariae,  of  diverse  sizes,  abound  on  the  surface. 

The  superomarginal  spines  stand  obliquely,  mostly  two  or  three 
on  a  plate.  They  are  longer  than  in  the  allied  species,  rather  stout, 
much  flattened,  and  often  a  little  grooved  on  one  side  distally,  but 
not  thickened  as  in  A.  anomala. 


Fi&  10. 


Atlosteriaj  forficulosa  Verrill.     No.   1183,  M.  C.  Z.     Spines;  a,  intra-ambulacral;  b,  outer- 
adambulacral;  c,  inferomarginal;   d,  superomarginal;  e,  dorsal.      X  20. 

The  inferomarginal  spines  are  very  similar  in  size  and  form ;  they 
stand  two  to  a  plate,  in  a  regular  double  row.  A  wide,  smooth 
channel  separates  the  two  series  of  marginal  spines,  and  a  similar 
channel  separates  the  adambulacral  series.  These  channels  have 
numerous  rather  large,  very  acute,  lanceolate  pedicellariae,  and 
many  small  papulae.  The  marginal  spines  all  bear  clusters  of  small 
minor  pedicellariae  on  the  outer  surface. 

The  adambulacral  spines  appear  crowded,  in  about  three  rows. 
They  mostly  stand  alternately,  one  and  two  to  a  plate.  The  inner  or 
furrow  series  stand  erect.  They  are  long,  slender,  very  acute,  and 
bear  clusters  of  major  pedicellariae  of  diverse  sizes,  but  mostly  very 
acute.  The  outer  spines  are  shorter,  stouter,  obtuse,  and  mostly 
bear  clusters  of  very  small  minor  pedicellariae  on  the  outer  side. 


196 


VERRILL 


The  larger  pedicellariae  of  the  lateral  channels  (o-r)  are  very 
acute-lanceolate  and  compressed.  The  larger  ones  of  the  dorsal 
surface  (a,  c,  d,  f)  are  similar  in  size  and  form,  but  some  are  stouter 
and  less  acute  (b,  h,  k),  and  there  are  with  them  many  of  smaller 
sizes  (t,  j,  I,  m,  n).  The  larger  and  longer  ones  measure 
i.oo  mm.  X  0.33  mm.;  i.oo  mm.  X  0.37  mm.;  the  stouter  ones, 
0.87  mm.  X  0.37  mm. ;  0.72  mm.  X  0.37  mm. 

The  type  was  from  Japan,  probably  Sagami  Bay  (No.  1183,  Mus. 
Comp.  Zool.). 


FIG.  ii. 


Allasteritu  forficulosa  Verrill.  No.  1183,  M.  C.  Z.  Pedicel  larise;  a-n,  major  pedicelUriae 
from  the  dorsal  surface;  o-r,  the  same  from  the  actinal  interradial  region  (axillary),  and  mar- 
ginal lanes.  X  32. 

The  Japanese  species  figured,  but  not  described,  by  Ives  (1891) 
appears  to  be  either  this  or  a  very  closely  related  species  of  the  same 
genus.  It  is  very  different  from  Liitken's  species. 

This  species,  though  not  known  from  the  American  side,  is  intro- 
duced here  for  comparison  with  A.  rathbum,  to  which  it  is  certainly 
pretty  nearly  allied.  It  has  fewer  adambulacral  spines,  larger  and 
longer  dorsals  and  marginals,  and  much  more  numerous,  larger, 
and  more  acute  major  pedicellariae.  It  is,  no  doubt,  related  to  Allas- 
terias  versicolor  (Sladen),  also  from  Japan,  as  mentioned  above. 

The  latter  has  far  less  numerous  dorsal  spines,  and  the  major  pedi- 
cellariae are  much  stouter  and  less  acute. 


SHALLOW-WATER   STARFISHES  197 

Subfamily  PYCNOPODIIN^E. 

Family  Pycnopodida  (pars)  STIMPSON,  op.  cit,  1861,  p.  261. 

Family  Pycnopodidee  FISHER,  op.  cit,  19066,  p.  136  (with  Rathbunaster  F.). 

Disk  broad  and  flat,  with  numerous  rays,  increasing  with  age  by 
the  interpolation  of  new  rays  by  budding  in  successive  pairs  bilater- 
ally, at  least  in  the  type. 

Dorsal  skeleton  feebly  developed,  the  ossicles  in  part  united  only 
by  thick  integument.  Spines  few,  scattered.  Pedicellariae  usually 
large  and  numerous,  of  two  kinds.  Adambulacral  plates  mostly 
monacanthid.  Marginal  plates  distinct.  Interactinal  plates  small  or 
lacking.  Body-cavity  without  the  circular  disco-brachial  partition, 
present  in  Heliasterinae. 

A  pair  of  gonads ;  a  pair  of  retractor  muscles ;  and  a  stomach-lobe 
for  each  ray.  Interbrachial  septa  feebly  developed.  Podia  large, 
mostly  in  four  rows  in  adults,  often  biserial  proximally  and  in  young. 
Ambulacral  plates  not  much  crowded,  sometimes  partially  alternate 
in  rays  that  are  bent  laterally. 

This  subfamily  includes,  among  living  species,  only  the  genera 
Pycnopodia  and  Rathbunaster,  each  with  a  single  species.  Rathbun- 
aster  californicus  Fisher  is  from  off  San  Diego,  California,  in  339 
fathoms. 

Some  fossil  starfishes  from  the  Devonian  closely  resemble  this 
group. 

Genus  Pycnopodia  Stimpson. 

Pycnopodia  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist,  vni,  p.  261, 1861.  Perrier, 
Revision,  Arch.  Zool.  Exper.,  iv,  p.  353,  1875.  Viguier,  op.  cit.,  vii,  p.  109, 
pi.  v,  figs,  ii,  12,  1878  (odontophore,  etc.).  A.  Agassiz,  North  American 
Starfishes,  p.  100,  pi.  xm,  1877  (structure  of  skeleton).  Sladen,  Voy. 
Challenger,  xxx,  pp.  xxxix,  560,  830,  1889.  Ritter  and  Crocker,  Proc. 
Wash.  Acad.  Sci.,  n,  p.  247,  plates  xiu,  xiv,  1900  (increase  of  number  of 
rays).  Loeb,  Publ.  Univ.  Calif.,  Physiol.,  11,  pp.  5-30,  1904  (experiments 
in  hybridization). 

Disk  large,  covered  with  a  thick,  soft  integument,  in  which  are 
few,  mostly  detached,  elliptical  ossicles,  some  of  which  bear  slender 
isolated  spines.  Clusters  of  minor  pedicellariae  are  also  scattered 
over  the  surface,  and  in  large  specimens  there  are  usually  many 
large,  scattered  major  pedicellariae.  Rays  variable  with  age,  up 
to  twenty  to  twenty-four  in  the  adult.  Their  increase  is  by  the  inter- 
polation of  new  rays,  usually  symmetrically  in  pairs,  between  the 
older  ones.  Superomarginal  and  inferomarginal  plates  are  ventral. 
They  form  two  regular  parallel  rows,  close  together,  united  by  the 


198  VERRILL 

broad  descending  apophyses  of  the  upper  plates,  with  large  inter- 
vening papular  areas.  The  upper  plates  are  large,  deeply  three-lobed, 
and  usually  bear  a  single  large  spine.  The  lower  plates  are  rather 
smaller,  less  strongly  lobed,  and  usually  bear  two  large  divergent 
spines.  Adambulacral  plates  are  crowded,  oblique,  bilobed  distally, 
and  each  bears  a  single  slender  spine.  Adoral  carina  is  long,  contain- 
ing twelve  or  more  pairs  of  contingent  ossicles.  Two  large  and  two 
smaller  peroral  spines.  Oral  area  large. 

This  genus  is  peculiar  to  the  west  coast  of  North  America.  It  con- 
tains only  a  single  species,  which  is  one  of  the  most  common  and 
most  characteristic  littoral  and  shallow-water  starfishes,  from  Puget 
Sound  to  Yakutat,  Alaska.  It  lives  among  stones  at  and  below 
low-tide  mark. 

PYCNOPODIA  HELIANTHOIDES  (Brandt)  Stimpson. 

Plate  xxix,  figure  i ;  plate  xxx ;  plate  xxxi,  figures  r,  2 ;  plate  LXXIH,  figure 
i;  plate  LXXIV,  figures  1-30  (young),  figure  6  (pedicellariae);  plate 
LXXXVIII,  figures  7-7<i  (details)  ;  text-figure  No.  2  (pedicellariae). 

""""  1^ 

Asterias  helianthoides  BRANDT,   Prod.   Desc.   Anim.,   Mertens,   p.   71,    1835. 

Stimpson,  Journ.  Bost.  Soc,  Nat.  Hist,  vi,  p.  89,  1857  (no  description). 
Pycnopodia  helianthoides  STIMPSON,  Proc.  Boston  Soc.  Nat  Hist,  rm,  p. 
261,  1861.  Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  pp.  324,  326,  327,  1867  (no 
description).  Perrier,  Revision,  Arch.  Zool.  Exper.,  nr,  p.  353,  1895  (no 
description).  A.  Agassiz,  N.  Amer.  Starfishes,  p.  100,  pi.  xm,  1877 
(structure).  Whiteaves,  Trans.  Royal  Soc.  Canada,  nr,  p.  n6,  1887  (dis- 
tribution). Ritter  and  Crocker,  Multiplication  of  Rays  and  Bilateral 
Symmetry,  Proc.  Wash.  Acad.  Sci.,  n,  pp.  247-274,  pis.  xm,  xiv,  figs,  1-13, 
1900.  Clark,  Proc.  Boston  Soc.  Nat.  Hist,  xxix,  p.  329  (no  description) . 

Disk  broad  and  soft.  Rays  in  the  adult  are  usually  from  eighteen 
to  twenty-four,  most  frequently  an  even  number,  but  those  with  odd 
numbers  are  not  rare.  I  have  seen  them  with  fifteen,  seventeen, 
nineteen  and  twenty-one  rays.  In  the  young  the  number  varies  from 
six  to  twelve  or  more.  The  increase  is  brought  about  by  the  budding 
in  of  successive  pairs  of  new  rays,  in  interradial  angles  situated 
symmetrically,  as  shown  by  Messrs.  Ritter  and  Crocker.1  Sometimes 
odd  new  rays  appear  in  other  places.  (See  also  our  pi.  xxxi.)  The 
radii  in  the  rather  small  dry  specimen  figured  on  our  pi.  xxix,  are 
75  mm.  and  182  mm. ;  ratio,  i :  2.42. 

The  rays  are  rather  slender  and  regularly  tapered,  covered  dorsally 
by  a  soft  skin,  which  bears,  chiefly  toward  the  ends  of  the  rays,  a  few 
scattered,  slender  spines  attached  to  slender,  detached  ossicles.  On 

1  Proc.  Wash.  Acad.  Sci.,  11,  p.  247,  pi.  xiv,  1900. 


SHALLOW-WATER   STARFISHES  199 

the  disk  some  of  the  ossicles  may  unite  into  a  feeble  network.  Thick 
clusters  of  minor  pedicellariae  are  also  scattered  over  the  dorsal  mem- 
brane. In  our  larger  specimens  (see  pi.  xxx),  there  are  also  many 
large  major  pedicellariae  scattered  over  the  dorsal  integument,  but 
they  are  sometimes  entirely  lacking  in  young  ones,  up  to  125  mm.  in 
diameter.  They  are  compressed,  long-ovate  in  outline,  obtuse  or  sub- 
acute,  not  dentate,  often  exceeding  the  spines  in  diameter  (text- 
fig.  2). 

The  dorsal  spines  are  sometimes  partly  short  and  obtuse,  scarcely 
longer  than  thick ;  most  are  long  and  slender,  cylindrical  or  tapered, 
rarely  subclavate.  They  are  often  striated  or  grooved  at  the  tip,  but 
more  commonly  are  regular  and  subacute.  They  are  more  numerous 
on  the  distal  part  of  the  rays,  where  they  become  very  slender  and 
bear  a  thick  wreath  of  minor  pedicellariae  at  about  mid-height. 
Those  on  the  disk  have  the  wreath  of  pedicellariae  around  the  base. 
The  superomarginal  spines  are  similar  to. the  dorsals,  but  a  little 
larger;  they  generally  stand  singly  on  alternate  plates,  and  bear 
thick  wreaths  of  pedicellariae.  The  inferomarginal  spines  are  longer 
and  stouter,  often  slightly  enlarged  at  both  ends,  and  mostly  flattened 
and  obtuse  or  subclavate  at  the  tip,  two  to  a  plate. 

The  adambulacral  plates  are  strongly  compressed ;  each  bears  a 
single,  very  slender,  slightly  tapered,  terete  spine.  Major  pedicel- 
lariae of  different  sizes  also  occur  on  the  adambulacral  plates,  with 
pedicels.  Some  of  these  are  even  larger  than  those  on  the  dorsal 
surface.  They  are  similar  in  form,  but  are  often  more  oblong- 
orate.  The  two  apical  peroral  spines  are  stouter  and  much  shorter 
than  the  adorals ;  the  side-spine  is  short  and  stout.  The  epiorals  are 
long  and  slender,  tapered,  longer  than  the  adorals.  The  adoral 
spines  are  decidedly  longer  than  those  more  distal. 

The  ambulacral  feet  are  in  four  regular  rows  in  half-grown  and 
mature  specimens,  except  proximally  and  distally,  where  they  are 
nearly  biserial.  In  quite  small  ones  they  are  in  two  nearly  regular 
rows.  , 

The  color,  in  life,  is  often  bright  red  to  reddish  brown,  but  varies 
from  yellow,  orange,  and  carmine  to  violet. 

Three  large  specimens  from  Puget  Sound  (coll.  Professor  Kincaid, 
Mus.  Comp.  Zool.,  No.  1908)  have  seventeen,  eighteen,  and  twenty 
rays,  respectively.  One  much  smaller  specimen,  from  the  same  place, 
has  seventeen  rays;  and  one  about  eight  inches  (200  mm.)  in 
diameter  has  eighteen  rays.  One  about  six  inches  (150  mm.)  in 
diameter  has  only  twelve  rays.  Thus  the  number  of  rays  does  not 


2OO  VERRILL 

depend  very  closely  on  the  size.  One  of  the  largest  of  this  lot  has 
the  radii  70  mm.  and  275  mm.  Some  young  specimens,  not  more 
than  30  mm.  in  diameter,  already  have  sixteen  rays;  while  others, 
200  mm.  in  diameter,  have  no  more. 

It  is  said,  on  good  authority,  to  become  over  four  feet  in  diameter. 
In  that  case,  it  is  probably  the  largest  starfish  known.  The  largest 
that  I  have  seen  are  over  two  feet  in  diameter.  Large  specimens 
are  hard  to  preserve,  and  for  that  reason  are  seldom  collected. 

TERATOLOGY. 

The  budding  in  of  new  rays  does  not  always  proceed  in  a  regular 
bilateral  manner  by  pairs.  I  have  studied  several  young  specimens 
in  which,  in  addition  to  the  regular  pairs,  one  or  two  new  rays  had 
appeared  irregularly  in  different  places.  If  one  odd  ray  appears  it 
gives  rise  to  an  odd  total  number  of  rays;  for  if  the  young  start 
with  six  rays  and  a  pair  of  new  rays  appears  each  time,  the  final 
number  must  be  even.  One  specimen  before  me,  eight  inches  in 
diameter,  has  nineteen  equal  and  regular  rays,  with  a  pair  of  small 
budding  rays  placed  regularly.  Such  an  individual  may  have  had 
seven  rays  at  first  instead  of  six.  In  most  cases  the  first  pair  of 
interpolated  rays  appears  either  side  of  a  single  ray,  with  five  rays 
on  the  other  side.  This  indicates  that  the  original  number  of  rays 
in  the  young  is  normally  six.  But  in  the  case  just  cited  there  are  five 
posterior  rays,  as  usual,  while  there  are  fourteen  (seven  pairs)  in 
front,  indicating  that  at  first  there  may  have  been  two  front  rays,  or 
seven  altogether.  This  may  be  the  manner  of  origin  of  some  speci- 
mens with  odd  numbers  of  rays.  But  I  have  seen  no  very  young 
ones  of  this  sort.  Should  one  bud  be  suppressed,  or  two  buds 
appear  together  on  one  side,  in  place  of  one,  the  final  result  would 
be  the  same. 

I  have  studied  very  young  ones  in  which  one  bud  is  suppressed 
(see  pi.  LXXV,  fig.  i).  One  young  example  (pi.  LXXV,  figs.  3,  30) 
has  the  madreporite  much  larger  than  usual.  It  has  an  oblong  disk 
and  eighteen  somewhat  unequal  rays.  It  looks  as  if  about  to  divide 
autotomously.  Its  disk  is  15  mm.  by  12  mm.  in  diameter.  The 
arrangement  of  the  rays  and  new  buds  in  this  specimen  is  peculiar 
and  anomalous.  Three  very  small  rays,  nearly  identical  in  size  and 
structure,  are  developing  simultaneously  at  three  places.  Two  of 
these  are  in  the  normal  position,  and  separated  by  seven  nearly 
equal  rays.  Then  follow,  on  one  side,  two  larger  rays  succeeded  by 
a  small,  extra,  budding  ray ;  on  the  other  side  no  such  bud  appears. 


SHALLOW-WATER   STARFISHES  2OI 

Between  the  extra  bud  and  the  normal  one,  on  the  other  side,  there 
are  six  normal,  equal  rays,  making  fifteen  normal  rays,  plus  three 
budding  ones. 

This  arrangement  does  not  conform  to  any  regular  rule.  The 
extra  bud  is  not  due  to  replacement  of  a  lost  ray,  for  it  originated 
like  the  others.  Before  it  appeared,  this  end  of  the  body  must  have 
had  eight  equal  rays,  which  would  have  been  abnormal,  for  five 
would  be  the  regular  number.  The  interpolation  of  the  new  extra 
bud  may  have  been  due  to  a  tendency  to  resume  an  odd  number  of 
rays  at  this  end,  which  is  the  usual  condition.  The  arrangement 
may  be  formulated  as  follows,  letting  the  figures  represent  normal 
rays  and  the  letters  budding  rays.  If  the  extra  bud  (6)  is  simply 
adventitious,  the  arrangement  would  be  thus : 

2    3    4    a    5    6          78 

i    • 

234a56b;8 

If  the  bud  (b)  be  for  regulation  purposes  it  would  stand  thus : 

234*5678 

i  9. 

234as6b8 

In  this  case  the  bud  would  pair  with  ray  7.  But  this  would  give  this 
end  of  the  starfish  nine  rays,  instead  of  the  normal  number,  five. 
(See  pi.  LXXV,  figs.  3,  30.)  Probably  autotomy  would  have  taken 
place  later,  followed  by  a  regulation  process  by  which  the  usual 
number  and  arrangement  of  rays  would  have  been  resumed  in  one 
or  both  parts. 

The  budding  in  of  rays  in  this  species  may  continue  till  late  in  life. 
A  specimen  studied  by  me,  nine  inches  in  diameter,  having  nineteen 
fully  developed  rays,  had  a  pair  of  small,  equal  budding  rays  in  the 
normal  position,  except  that  there  were  fourteen  rays  on  the  budded 
side,  instead  of  an  odd  number.  There  were  five,  as  usual,  on  the 
other  side. 

A  careful  study  of  the  living  young  of  this  species  should  be 
made,  where  it  is  abundant,  to  ascertain  more  fully  its  complete 
history  during  growth. 

In  some  cases  the  ambulacral  plates  are  not  strictly  opposite,  and 
may  even  appear  to  be  alternate,  when  the  rays  are  strongly  bent 
sidewise,  owing  to  their  loose  articulations.  In  a  few  cases  the 
number  of  these  plates  is  not  the  same  on  the  two  sides  of  the  groove, 
for  a  short  distance ;  apparently  due  to  the  irregular  interpolation  of 
an  extra  plate  on  one  side  only,  here  and  there.  When  this  occurs 


2O2  VERRILL 

the  plates  must  become  alternate,  locally.  This  recalls  the  regular 
alternation  of  these  plates  in  many  paleozoic  fossil  starfishes. 

The  species  has  an  extensive  range,  from  Monterey  and  Tomales 
Bay,  California,  to  Dutch  Harbor  and  Yakutat,  Alaska.  According 
to  Professor  Ritter,  it  was  very  abundant  at  the  latter  place  on  a 
rocky  reef  exposed  at  low  tide,  while  the  young  were  found  adhering 
to  Laminaria.  It  is  also  very  abundant  at  Sitka  and  in  Puget 
Sound. 

Gulf  of  Georgia  (Whiteaves)  ;  Gulf  of  Singio  and  California 
(Perrier) ;  Puget  Sound  and  Tomales  Bay  (Stimpson)  ;  Port 
Townsend,  Puget  Sound,  abundant  (Clark).  It  occurs  at  low  tide 
and  in  shallow  water,  on  rocks. 

I  have  also  examined  specimens  from  near  Monterey,  California ; 
Tomales  Bay;  Victoria;  Puget  Sound  (Kincaid)  ;  Vancouver  Island 
(Canadian  Geological  Survey)  ;  Queen  Charlotte  Islands,  etc. 

It  was  collected  by  the  Harriman  Expedition  at  Sitka,  Dutch 
Harbor,  and  Yakutat,  Alaska. 

Family  PEDICELLASTERID 2E  Perrier  (pars). 
Ptdicellosterida  SLADEN,  op.  cit,  1859,  p.  556. 

As  adopted  by  Sladen,  this  family  included  only  the  genus  Pedicel- 
laster.  It  was  distinguished  from  Asteriidae  only  by  the  arrangement 
of  the  podia  or  "  sucker-feet "  in  two  rows,  instead  of  four.  More 
recently  Perrier  has  added  the  genus  Gastraster,  which  has  some  of 
the  podia  in  four  rows. 

Genus  Pcdiccllaster. 
Type,  P.  typicus  M.  Sars. 

Pedicellaster  M.  SARS,  Oversigt  over  Norges  Echinod.,  p.  77,  1861.  Sladen, 
op.  cit.,  1859,  p.  557. 

Small,  delicate  starfishes  with  a  reticulated  dorsal  skeleton,  bearing 
slender  spinules ;  two  rows  of  similar  marginal  spines.  Adambulacral 
spines  usually  diplacanthid,  rarely  triplacanthid,  or  partially  so. 
Minor  pedicellariae  on  dermis,  similar  to  those  of  Leptasterias. 
Major  pedicellariae  often  lacking.  Rays  five  or  six ;  sometimes  eight. 

PEDICELLASTER  TYPICUS  M.  Sars. 

Pedicellaster  typicus  M.  SARS,  op.  cit.,  1861,  p.  77,  pi.  ix,  figs.  9-17,  pi.  x,  figs, 
i-io.  Danielssen  and  Koren,  Norske  Nordhavs-Exped.,  Zool.,  xi,  Aster- 
oidea,  pp.  36-40,  1884.  Sladen,  op.  cit.,  1889,  p.  557. 


SHALLOW-WATER  STARFISHES  2OJ 

Pedicellaster  palaocrystallus  SLADEN,  Ann.  and  Mag.  Nat.  Hist.,  ser.  5,  rol.  v, 
p.  216,  1880.  Duncan  and  Sladen,  Mem.  Echinod.  Arctic  Sea,  W.  of  Green- 
land, p.  34,  pi.  n,  figs.  22-26,  1881. 

A  single  small  specimen  of  this  delicate,  five-rayed  species  (diam- 
eter, 1  6  mm.)  occurred  in  a  lot  of  other  small  starfishes  sent  by  the 
National  Museum,  No.  6123.  It  was  from  the  Arctic  coast  of  Alaska, 
near  Icy  Cape,  in  10  to  15  fathoms,  mud  and  sand  (Smith  coll.), 
through  W.  H.  Dall,  1874. 

This  species  is,  therefore,  circumpolar.  It  has  been  taken  off 
Greenland,  Grinnell  Land,  Cape  Fraser,  Nova  Zembla,  Beeren  Island, 
etc.,  and  extends  southward  to  the  Norwegian  coast  and  to  the  Gulf 
of  St.  Lawrence.  Its  bathymetrical  range  is  from  15  to  620  fathoms. 

Order  SPINULOSA  Perrier. 

Stellerida  Spinulosee  PERKIER,  Nouv.  Arch.  Mus.  Hist.  Nat,  vi,  p.  154,  1884. 
Spinulosa  PERKIER,  Exped.  Trav.  et  Talism.,  pp.  27,  138,  1894. 
Phanerozonia  and  Cryptozonia  (pars)  SLADEN,  Voy.  Chall.,  xxx,  p.  xxiii,  1889. 
Spinulosa  FISHER,  op.  cit,  igiib,  p.  251  (table  of  known  families). 


This  order,  established  by  Perrier,  includes  cryptozonate  (rarely 
subphanerozonate)  starfishes  which  usually  have  but  two  rows  of 
ambulacral  feet  (four  in  Diplopteraster)  ;  ambulacra!  plates  not 
crowded;  and  generally  a  reticulated  or  imbricated  dorsal  skeleton, 
but  rarely,  if  ever,  true  paxillae.  The  ossicles  are  sometimes  isolated 
or  vestigial  in  deep-sea  genera.  The  pedicellariae  are  generally  lack- 
ing. When  present,  they  are  neither  forcipulate  nor  f  orficulate.  They 
may  be  pincer-like,  with  two  or  more  simple  valves,  and  are  always 
dermal.  The  marginal  plates  are  nearly  always  small,  and  sometimes 
indistinct.  The  lower  ones  are  usually  the  larger.  They  never 
form  a  thick,  rigid  margin;  the  margin  may  be  acute  or  rounded. 
Papulae  may  occur  both  on  the  dorsal  and  on  the  actinal  surfaces, 
but  they  are  restricted  to  the  dorsal  surface  in  most  families. 

The  odontophores  are  adambulacral  of  various  forms.  The  ambu- 
lacral grooves  are  usually  narrow.  The  ambulacral  feet  always  have 
a  terminal  sucker.  The  ambulacral  ampullae  may  be  single  or 
double. 

Perhaps  the  structure  of  the  pedicellariae  is  the  most  positive  char- 
acter for  separating  certain  genera  of  this  group  from  some  of  the 
aberrant  Forcipulosa. 

A  large  number  of  the  genera  do  not  have  free-swimming  larvae, 
but  are  known  to  carry  their  eggs  and  larvae  attached  about  the 
mouth,  or  otherwise,  or  in  a  marsupial  pouch  (gonocodium)  ,  until 


204  VERRILL 

they  become  true  starfishes,  large  enough  to  care  for  themselves.1 
This  habit  is  usually  associated  with  the  lack  of  pedicellariae.  It  is 
conducive  to  the  formation  of  local  varieties.  The  order  may  be 
conveniently  divided  into  two  suborders: 

I.  AVELATA  Verrill. 
II.  VELATA  Perrier. 

Suborder  AL VELATA  Verrill,  nom.  nov. 

This  includes  the  more  typical  forms  in  which  there  is  no  dorsal, 
tent-like  marsupial  chamber  for  the  protection  of  the  eggs  and  young. 
The  spines  are  either  all  free,  or  partially  or  wholly  webbed  together 
into  groups,  as  in  Solasteridae,  in  which  the  groups  of  adambulacral 
spines  form  transverse  combs  completely  webbed  together,  and  the 
dorsal  spinules  are  partially  so. 

It  includes  the  following  families,  represented  on  the  Northwest 
coast  in  shallow  water : 

Family  ECHINASTERID&. 
Family  SOLASTERIDJE. 
Family  MITHRODIWJE. 
Family  ASTERINID&. 

Also  the  following  extraliniital  or  deep-sea  families,  besides  other 
smaller  groups : 

GANERIID&,  Patagonian ;  Antarctic. 
ACANTHASTERIDJE,  Tropical;  Panamic. 
PYTHONASTERIDJE,  Deep  Sea. 
KORETHRASTERIDJE,  Deep  Sea. 
MYXASTERID&,  Deep  Sea. 

Suborder  VELATA  Perrier. 

This  group  includes  only  the  family  Pterasteridae.  It  is  remark- 
able for  having  most  of  the  spines  webbed  together  in  clusters  and 
for  the  remarkable  development  of  a  dorsal  membrane,  more  or 
less  completely  uniting  the  parapaxillae  together,  forming  a  dorsal 
marsupial  pouch  or  gonocodium  in  which  the  eggs  and  young  are 
carried. 

M.  Perrier,  in  his  later  works,  has  separated  the  Pterasteridae  as 
an  order  (Velata).  This  may  be  overestimating  the  importance  of 

1  This  is  the  case  with  the  genera  Henricia,  Solaster,  Pteraster,  Hymenaster, 
and  others. 


SHALLOW-WATER   STARFISHES  2O5 

the  peculiar  characters  of  the  group.  The  Velata  must,  however,  be 
regarded  as  a  highly  specialized  group,  or  suborder,  differing  in 
many  respects  from  all  other  starfishes.  The  Pythonasteridae  seem 
to  me  more  nearly  allied  to  Solasteridae. 

The  family  Valvasteridae,  referred  to  Spinulosa  by  Fisher,  seems 
to  me  to  belong  rather  to  Valvulosa,  among  the  Phanerozona.  It  is 
not  represented  on  the  west  coast 

Family  ECHINASTERID&  Verrill  (restricted). 

Echinasteridee  VERRILL  (pars),  Trans.  Conn.  Acad.  Sci.,  i,  p.  343,  1867.  Per- 
rier  (pars),  Revis.  Stell.,  Arch.  Zool.  iv,  pp.  299,  358,  1875. 

Echiniasterinee  VIGUIER,  Arch.  Zool.  Exper.  et  Gen.,  vii,  p.  123,  1878  (struc- 
ture). 

Echtnasteridee  SLADEN  (Pars),  Voy.  Chall.,  xxx,  p.  535,  1889.  Bell,  Catal. 
Echinod.,  pp.  23,  95,  1890.  Perrier,  Etoiles  de  met,  Nouv.  Arch.  Mus. 
Nat.  Hist.,  vi,  p.  164,  1884;  Exp.  Trav.  et  Talism.,  pp.  28,  141,  1894.  Lud- 
wig,  Fauna  Arctica,  i,  p.  472,  1900.  Fisher,  op.  cit.,  19116,  p.  258  (analyt- 
ical table  of  known  genera,  p.  259). 

Dorsal  skeleton  usually  formed  of  small,  sometimes  overlapping, 
reticulated  or  areolated  ossicles;  sometimes  they  form  longitudinal 
radial  rows.  Median  dorsal  row  is  often  indistinct.  Upper  mar- 
ginals small,  often  not  specialized.  Inferomarginals  usually  distinct. 
One  or  more  interactinal  rows  are  usually  present.  Dorsal  and 
marginal  spines  may  be  large  and  isolated  or  minute  and  clustered, 
but  not  truly  paxilliform.  They  are  often  covered  with  a  distinct, 
thick,  dermal  membrane,  which  may  contain  calcareous  granules. 

The  ambulacra!  ampullae  are  usually  single,  one  to  each  tube-foot. 

Ambulacral  grooves  narrow.  Adambulacral  spines  small,  two  or 
several  to  a  plate,  either  in  longitudinal  or  in  transverse  groups.  No 
pedicellariae  known. 

The  most  prominent  genera  are  Echinaster  and  Henricia,  both  of 
which  occur  on  the  Northwestern  coast. 

Genus  Poraniopsis  Perrier. 

Poraniopsis  PERRIER,  op.  cit,  1891,  p.  105,  pi.  x,  figs.  20,  2&  (type  P.  tchinaster 

Perrier).     Fisher,  1910,  p.  568;  I9H&,  p.  260. 
LahilUa  DE  LORIOL,  1904,  p.  32,  pi.  in,  figs,  l-lg  (preoccupied). 
Alexandraster   LUDWIG,    1905,   p.    210,   pi.    xv,   figs.   79,   80;   pi.    xxxi,   figs. 

.    181-183 ;  pl-  xxxii,  fig.  184. 
Ortmannia  DE  LORIOL,  op.  cit.,  1906,  p.  78. 

Form  is  five-rayed,  stellate.  Surface  covered  with  a  thick  skin. 
Abactinal  ossicles  spinose,  slender,  forming  an  open  reticulation  with 


206  VERRILL 

wide  papular  areas  and  numerous  papulae.  Papulae  lacking  on  actinal 
side,  present  between  marginals.  Adambulacral  plates  bear  two 
spines;  none  in  the  furrow.  Pedicellariae  unknown.  Madreporite 
an  independent  plate. 

PORANIOPSIS  INFLATA  Fisher. 

Alexaridroster  inflates  FISHER,  op.  cit.,  1906,  p.  300. 

Poraniopsis  inflate  FISHER,  191  it,  p.  261,  pi.  LVIII,  figs.  7,  70;  pi.  LXIII,  figs.  I, 
2;  pi.  cxn,  fig.  i. 

Form  stellate,  with  five  short,  thick,  tapered  rays.  Radii  of  the 
type,  23  mm.  and  60  mm. ;  ratio,  1 : 2.6.  Dorsal  spines  large,  3  mm. 
to  6  mm.  high,  acute,  usually  one  at  each  angle  of  the  large  papular 
areas,  forming  altogether  about  seven  imperfect  rows  in  the  type, 
not  including  the  inferomarginals  and  those  usually  present  on  the 
interactinal  plates.  Adambulacral  spines  two  to  a  plate,  grooved, 
as  are  usually,  also,  the  interactinal  spines,  when  present.  The 
plates  and  spines  are  covered  with  thick  membrane. 

Recorded  by  Fisher  from  seventeen  localities,  in  26  to  159  fathoms, 
from  Oregon  to  San  Diego,  California. 

It  was  not  in  the  collections  received  from  farther  north.  Dr. 
Fisher  described  another  form,  from  deeper  water  (334  to  600 
fathoms),  as  P.  inflata  fie.vibilis  Fisher. 

Genus  Echinaster  Miiller  and  Troschel ;  includes  Othilia  Gray. 

Stellonia  (pars)  NARDO,  Oken's  Isis,  p.  716,  1834.    Agassiz,  Prod.,  p.  191,  1835. 

Echinaster  (pars)  MULLER  and  TROSCHEL,  Monatsber.,  Berlin  Wiss.  Akad., 
April,  1840,  p.  102;  Wieg.  Arch.,  1840  (no*  Gray);  Syst.  Aster.,  p.  22, 
1842  (pars). 

Othilia  GRAY,  Ann.  Mag.  Nat.  Hist.,  vi,  p.  281,  Dec.,  1840;  Synopsis  Starfishes, 
p.  12,  1866.  Fisher,  191  ib,  p.  260. 

Echinaster  PERRIER,  Revis.  Stell.  du  Mus.,  Arch.  Zool.  Exper.,  iv,  p.  364,  1875. 
A.  Agassiz,  North  Amer.  Starfishes,  p.  97,  pi.  x,  1877  (structure  of  skele- 
ton). Viguier,  Squelette  Stell.,  Arch.  Zool.  Exper.,  vii,  p.  123,  pi.  vii,  figs. 
1-7,  1878  (structure  of  skeleton).  Sladen,  Voy.  Chall.,  Zool.,  xxx,  p.  553, 
1889.  Perrier,  Exp.  Trav.  et  Talism.,  p.  146,  1894. 

Disk  rather  small.  Rays  five,  of  moderate  length,  rounded. 
Larger  dorsal  and  marginal  ossicles  strong,  lobate,  convex  in  the 
middle,  and  having  a  central  mammilla  and  pit  for  the  attachment  of 
the  usually  solitary  spine.  The  larger  ossicles  are  united  into  reticu- 
lations by  small,  rounded,  connective  ossicles,  leaving  large  papular 
areas;  these  may  be  intermarginal  and  often,  also,  interactinal. 


SHALLOW-WATER   STARFISHES  20? 

The  whole  surface  is  covered  in  life  with  a  rather  thick,  soft 
integument,  containing  small  granules  on  the  spines.  Dorsal  spines 
mostly  isolated,  sometimes  in  small  groups,  conical,  acute. 

Upper  and  lower  marginal  plates  small,  not  very  distinct,  with 
spines  like  the  dorsals.  Adambulacral  spines  in  a  small  transverse 
row  of  two  to  four ;  the  inner  ones  are  smaller,  inserted  in  the  groove, 
just  below  the  margin. 

The  genus  Othilia  Gray,  December,  1840,  appears  to  be  a  syno- 
nym of  Echinaster  Miiller  and  Troschel,  April,  1840.  The  type  of 
the  latter  was  E.  spinosus  =  Asterias  echinophcra  Lam.,  both  by 
virtual  tautology  and  by  designation.  (See  foot-note  in  Muller  and 
Troschel,  Syst.  Ast.,  p.  22,  1842.)  Echinaster  of  Gray,  1866,  is 
Acanthaster  of  later  writers.  A.  echinophora  Lam.  appears  to  be 
identical  with  A.  spinosa  Retz.,  1783,  the  type  of  Othilia.  The  name 
spinosus  cannot  be  used  for  any  species,  because  it  was  preoccupied 
by  A.  spinosa  Muller,  1777. 

Fisher  (op.  cit,  191  ib,  p.  260)  separated  Othilia  from  Echinaster 
because  it  has  intermarginal  and  sometimes,  also,  actinal  papulae.  The 
type  of  Echinaster  and  also  E.  sentus  (Say)  have  interactinal 
papulae,  as  do  the  other  American  species. 

Echinaster  of  Fisher  is  probably  Rhopia  Gray,  but  he  does  not 
give  the  type  in  his  table.  The  type  of  the  latter  was  R.  seposita  of 
the  Mediterranean.  Although  that  was  the  first  species  mentioned 
under  Echinaster  by  Muller  and  Troschel,  April,  1840,  their  later 
designation  of  E.  spinosus,  as  the  type,  seems  to  settle  the  question  of 
the  type  of  Echinaster. 

ECHINASTER  (OTHILIA)  ROBUSTUS  Verrill,  sp.  nov. 

Disk  rather  large.  Rays  five,  large,  stout,  rather  short,  obtuse. 
Radii,  15  mm.  and  42  mm. ;  ratio,  1 : 3.5.  The  disk  is  flattened  in 
drying  and  doubtless  measures  more  than  it  would  in  life. 

The  dorsal  spines  are  large,  conical,  acute;  nearly  equal,  isolated 
and  pretty  regularly  spaced,  the  distances  between  being  about 
3  mm.  to  4  mm. ;  height  of  the  spines,  2  mm.  to  3  mm.  The  papular 
pores  are  numerous  and  have  large  angular  or  rounded  papular 
areas  between.  The  dorsal  and  upper  marginal  spines  form  about 
seven  rows,  but  they  stand  nearly  in  quincunx,  so  that  the  rows  are 
not  very  distinct. 

On  the  actinal  surface  there  is  one  row  of  ossicles  bearing  spines, 
the  inferomarginals,  in  contact  with  the  adambulacrals.  In  this  row, 
one,  two,  three,  and  sometimes  four,  stand  on  one  plate.  Sometimes 


208  VERRILL 

there  is  an  additional  imperfect  interpolated  row  below  the  supero- 
marginals  distally.  The  ventral  spines  are  rather  smaller  than  the 
dorsals. 

The  adambulacral  spines  mostly  form  a  transverse  group  of  three 
on  the  distal  plates ;  but  proximally  often  one  and  two  on  alternate 
plates.  The  inner  or  furrow  spine  is  much  the  smallest,  and 
stands  just  within  the  margin;  it  is  tapered,  acute,  and  about  half 
as  long  as  the  outer  ones.  The  latter  are  stouter  and  acute,  but 
smaller  than  the  marginals.  When  three  are  present,  the  inter- 
mediate one  is  intermediate  in  size.  The  outer  ones  are  somewhat 
webbed  together  at  base. 

Interactinal  plates  and  spines  are  absent.  The  inferomarginals  are 
united  directly  to  the  adambulacrals,  the  latter  being  transversely 
oblong  at  the  surface.  Two  or  three  short,  irregular  rows  of  inter- 
marginal  ossicles  are  interpolated  between  the  upper  and  lower  mar- 
ginals at  the  bases  of  the  rays.  The  large  spiniferous  dorsal  ossicles 
are  thick,  convex  and  lobulated.  The  smaller  connective  ossicles 
are  rounded  and  elliptical,  thick,  or  biscuit-shaped;  and  many  of 
them  have  a  naked,  glistening,  round,  central  area,  covered  with 
very  uniform  minute  punctulations.  Similar  punctate  ossicles  occur 
on  the  lateral  and  ventral  surfaces,  between  the  spines. 

The  two  apical  oral  spines,  and  the  pair  of  epioral  spines,  are  about 
equal  and  similar  to  the  adambulacrals,  but  not  so  large.  The 
lateral  adoral  spines  are  small  and  strongly  divergent.  A  few 
solitary  papulae  stand  between  the  inferomarginal  and  adambulacral 
plates.  Ambulacral  feet  and  pores  are  large.  Madreporic  plate 
small,  prominent,  with  few  rough  gyri. 

The  type  is  from  Sooke,  Vancouver  Island  (Canadian  Geological 
Survey,  1893),  received  through  Mr.  J.  F.  Whiteaves. 

ECHINASTER  TENUISPINUS  Verrill. 
Plate  cvn,  figure  2  (type). 

Echinaster  tenuispinus  VERRILL,  Trans.  Conn.  Acad.  Sci.,  i,  pp.  577,  594,  1867. 
Ives,  Proc.  Acad.  Nat  Sci.  Philadelphia,  1889,  p.  171;  List  of  Echinod., 
Cabinet  of  Stearns,  p.  ii,  1891.  Sladen,  Voy.  Chall.,  xxx,  pp.  554,  812, 
1889.  ??  Clark,  Proc.  Boston  Soc.  Nat.  Hist,  xxix,  p.  319,  1901. 

A  large  five-rayed  species  with  long,  tapering  rays,  swollen  at 
base,  but  becoming  slender  toward  the  tips.  Radii,  14  mm.  and 
91  mm. ;  ratio,  1 : 6.  This  specimen  is  one  of  the  original  types. 

Dorsal  spines  unusually  numerous  for  this  genus,  but  mostly 
isolated,  rather  small,  usually  I  mm.  to  2  mm.  long,  slender,  acute ; 


SHALLOW-WATER   STARFISHES  2O9 

they  are  scattered  over  the  surface,  somwehat  in  quincunx,  but  in 
some  parts  seem  to  belong  to  about  twelve  to  sixteen  rows. 

The  dorsal  ossicles  are  strong,  deeply  lobulate,  and  openly  reticu- 
late, leaving  many  papular  areas,  with  numerous  small  papulae.  The 
madreporic  plate  in  this  specimen  is  double,  the  two  parts  nearly 
equal,  rounded,  and  a  little  separated,  covered  with  small  spinules. 
The  superomarginal  plates  are  scarcely  distinguishable  from  the 
dorsal  ossicles,  except  distally ;  they  mostly  bear  one  or  two  conical 
spines.  The  inferomarginals  are  a  little  larger,  strongly  lobed,  and 
often  bear  two  spines  not  distinctly  larger  than  the  dorsals.  A  row 
of  small  interactinal  ossicles,  part  of  them  bearing  small  spines,  is 
interpolated  proximally.  The  adambulacral  spines  stand  in  a  trans- 
verse row  of  three,  or  sometimes  four,  on  each  plate,  besides  a 
small,  short  one  (rarely  two)  within  the  margin  of  the  groove.  They 
are  subequal,  round,  a  little  tapered,  blunt,  and  decidedly  longer  than 
the  outer  spines.  The  one  next  the  margin  of  the  groove  is  usually, 
but  not  always,  the  longest ;  the  outer  one  is  the  shortest. 

This  species  occurs  commonly  from  Panama  to  Lower  California. 
It  is  said  to  occur  also  at  San  Diego,  California. 

The  types  were  from  La  Paz,  Gulf  of  California ;  San  Diego  (Ives, 
1891) ;  Bay  of  Pichilinque  (Ives,  1889) ;  Monterey,  California 
(Ives,  1889).  The  latter  is  probably  an  error;  it  needs  confirmation. 

Mr.  Clark,  op.  cit.,  1901,  mentions  a  specimen  from  Puget  Sound, 
which  he  refers  doubtfully  to  this  species,  as  having  the  rays  "  long 
(52  mm.  to  62  mm.)  and  very  slender,  and  though  rough,  they  are 
not  spiny."  This  must  be  a  different  species  or  genus  if  it  has  no 
obvious  spines. 

Perhaps  the  locality  label  was  erroneous,  as  was  the  case  with 
several  other  species  in  the  same  collection.  It  was  not  to  be  found 
when  I  studied  the  collection,  at  Columbia  University. 

Genus  Henricia  Gray. 

Henricia  GRAY,  Ann.  Mag.  Nat.  Hist.,  vi,  p.  184,  November,  1840;  Synopsis 
Starfishes,  p.  5,  1866.  Bell,  Ann.  and  Mag.  Nat.  Hist.,  vr,  p.  473,  1890; 
Catal.  British  Echinod.,  p.  95,  1892.  Fisher,  191  ib,  p.  266. 

Rhopia  (pars)  GRAY,  op.  cit,  1866,  p.  12. 

Cribella  FORBES,  Brit.  Starfishes,  p.  100,  1841,  or  December,  1840. 

Echinaster  (pars)  MULLER  and  TROSCHEL,  April,  1840;  Syst.  Aster.,  p.  22, 
1842. 

Linckia  FORBES  (non  Nardo),  Mem.  Wern.  Soc.,  vm,  p.  120,  1839. 

IS 


2IO  VERRILL 

Cribrella  FORBES,  Brit.   Starfishes    (as  Cribella),   1841    (non  Agassiz).     L. 

Agassiz  (Pars),  Prod.,  in  Mem.  Soc.  Sci.  Nat.  Neuchatel,  i,  p.  191,  1835. 

Liitken,  Gronl.  Echinod.,  p.  30,  1857.    Norman,  Ann.  and  Mag.  Nat.  Hist., 

xv,  p.  124,  1865.    Verrill,  Proc.  Boston  Soc.  Nat  Hist.,  x,  p.  345,  1866. 

Perrier,  Revis.  Stell.  Mus.,  Arch.  Zool.  Exper.,  iv,  p.  373,  1875.    A.  Agassiz, 

North  Amer.  Starfishes,  p.  113,  pi.  xvni,  1877  (structure  of  skeleton). 

Viguier,   Squelette  des   Stell.,  Arch.  Zool.  Exper.,  vu,  p.   126,  pi.  vn. 

figs.  8-15,  1878  (odontophore).     Sladen,  Voy.  Chall.,  xxx,  p.  540,  1889. 

Perrier,  Exp.  Trav.  et  Talism.,  p.  143,  1894.    Ludwig,  Fauna  Arctica,  p. 

473- 

Form  usually  neatly  stellate  with  slender  terete  rays;  sometimes 
with  short  rays. 

Dorsal  ossicles  small,  convex,  closely  united  or  overlapping  at  the 
edges,  and  arranged  in  a  reticulated  or  areolated  pattern,  leaving 
small  papular  areas,  carrying  few,  often  single,  papulae,  and  not  cov- 
ered with  a  thick  integument.  Dorsal  spines  minute  and  nearly  uni- 
form, crowded  in  divergent  clusters  on  the  convex  ossicles,  both  on 
the  dorsal  and  lateral  surfaces.  Adambulacral  spines  in  transverse, 
usually  double  rows,  or  in  multiple  clusters  of  small  graded  spines 
on  the  outer  side  of  the  plates ;  the  inner  spine  is  compressed  and 
inserted  within  the  margin  of  the  groove  as  a  furrow-spine;  some- 
times two  are  present  in  the  grooves.1 

Superomarginals  small,  often  indistinct  and  like  the  dorsals. 
Inferomarginal  plates  small,  but  distinct,  often  larger  than  the 
superomarginals,  contingent  with  the  latter  distally,  but  separated 
proximally  by  one,  two,  or  more  rows  of  interpolated  marginal 
ossicles,  varying  in  number  and  extent,  and  thus  causing  great 
variations  in  the  thickness  and  taper  of  the  rays.  Peractinals  and 
inferomarginals  similar  and  closely  joined,  convex,  and  covered  with 
small  spinules,  the  rows  separated  by  papular  pores;  the  papulae 
may  occur  also  between  the  peractinals  and  adambulacrals. 

Pedicellariae  have  not  been  observed  in  any  species.  The  eggs  and 
young  are  carried  under  the  oral  region. 

The  madreporite  is  generally  covered  with  spinules  like  those  on 
the  other  dorsal  plates. 

The  West  Indian  species,  H.  antillarum  (Perrier,  as  Cribrella), 
has  six  rays  and  two  madreporites.  It  appears  to  be  autotomous. 

1 A  new  genus,  HENRICIDES,  is  now  proposed  for  Henricia  heteractis  Clark 
(op.  cit,  1909,  p.  530,  pi.  XLIX,  figs.  I,  2),  from  Australia, 

It  has  six  or  seven  rays,  multiple  madreporites,  and  is  probably  autotomous. 
Dorsally  it  is  like  Henricia.  It  has  no  furrow-spines;  the  outer  side  of  the 
adambulacral  plates  bears  a  marginal  row  or  comb  of  five  or  six  spines  pro- 
jecting over  the  furrow,  and  back  of  these  a  group  of  divergent  spines. 


SHALLOW- WATER    STARFISHES  211 

This  genus  differs  from  Echinaster  in  the  absence  of  thick  external 
integument;  in  the  covering  of  minute  spinules;  and  in  the  nearly 
uniform  small  ossicles  of  the  dorsal  and  lateral  skeleton,  the  con- 
nective ossicles  being  almost  indistinguishable  from  the  primary 
series.  There  are  no  large  mammillate,  spiniferous  ossicles,  nor  any 
intermediate  naked  ones,  above  or  below.  The  papular  areas  are 
much  smaller  and  the  papulae  stand  singly  or  in  small  groups. 

In  the  species  that  I  have  studied  there  is  but  one  well  developed 
row  of  interactinal  plates,  sometimes  as  large  as  the  inferomarginals 
proximally,  though  generally  much  smaller,  and  often  bearing  finer 
spines.  This  row  of  peractinals  may  extend  merely  to  the  middle  of 
the  ray,  but  it  often  disappears  only  close  to  the  tip.  Its  ossicles  are 
closely  joined  to  the  outer  side  of  the  adambulacrals,  which  they 
about  equal  in  length,  and  consequently  in  number,  but  in  some 
species  there  are  intervening  papulae.  In  some  cases  a  second  short 
row  of  synactinals  is  developed  proximally,  in  large  adult  specimens. 
The  adambulacral  plates  are  transversely  oblong. 

According  to  the  accepted  rules  of  priority  it  seems  necessary 
to  adopt  Henricia  instead  of  Cribrella  as  the  name  of  this  genus. 

The  generic  name  Cribrella  was  proposed  tentatively  by  Agassiz 
(op.  cit.,  p.  191,  1835),  as  a  substitute  for  Linckia  Nardo  (Isis, 
1834),  evidently  because  the  latter  had  been  previously  used  in 
botany.  Agassiz  did  not  then  actually  use  it  as  the  name  of  the 
genus,  but  retained  Linckia.  The  species  that  he  cited  were 
L.  variolata  N.,  L.  typns  N.,  and  L.  franciscus  N.  These  belong  to 
the  Ophidiasteridae,  and  to  Linckia  as  now  restricted.  Therefore  the 
generic  name,  Cribrella,  if  used  at  all,  must  be  considered  as  dating 
only  from  Forbes,  who  published  it  in  1841'  (or  December,  1840,  at 
the  earliest),  spelling  it  " Cribella"  evidently  by  a  typographical 
error,  and  applied  it  wrongly  to  the  present  genus. 

J.  E.  Gray,  however,  as  shown  by  Bell,1  had  published  a  valid  name, 
Henricia,  for  the  genus  at  a  slightly  earlier  date.  (Op.  cit.,  Novem- 
ber, 1840.) 

Therefore,  if  the  monthly  part  of  Forbes's  work,  including  Cri- 
bella, was  published  December  i,  1840,  as  Bell  states,  the  name  Hen- 
ricia evidently  has  one  month  of  priority  and  should  be  adopted. 
His  work,  as  a  whole,  is  dated  1841. 

'Op.  cit.,  vi,  p.  473,  1890.  Bell  there  states,  on  the  authority  of  the  pub- 
lishers, that  Forbes's  "British  Starfishes"  was  published  in  six  monthly 
numbers,  from  October,  1840,  to  March,  1841. 


212  VERRILL 

Moreover,  as  used  by  Agassiz,  1835,  it  was  a  synonym  of  Linckia, 
and  if  the  latter  name  be  retained,  it  must  be  rejected  on  that 
account  also. 

In  the  system  of  Agassiz,  1835,  this  genus  would  have  been 
included  under  Stellonia  Ag. ;  together  with  Echinaster,  Solaster, 
Heliaster,  Asterias,  etc. 

This  genus  is  remarkably  well  represented  on  the  North  Pacific 
coast.  Professor  Fisher  (ignb)  has  described  numerous  species 
and  named  varieties  or  subspecies  from  the  Northwest  coast,  as  well 
as  various  additional  minor  varietal  forms,  not  named. 

The  genus  is  a  very  difficult  one  and  the  number  of  forms  on  the 
Northwest  coast  is  surprisingly  great,  as  compared  with  those  of  the 
North  Atlantic,  where  but  one  species  is  usually  recognized 
(H.  sanguinolenta) ,  at  least  on  the  American  side.1 

The  latter,  from  our  coast,  though  variable  to  a  considerable  extent 
in  form  and  spinulation,  presents  no  such  extensive  variations  as  it 
and  other  species  show  on  the  Pacific  side.  I  have  personally 
examined  more  than  ten  thousand  specimens  of  the  Atlantic  san- 
guinolenta, from  a  great  variety  of  localities,  and  at  least  several 
hundreds  of  the  allied  North  Pacific  forms,  and  can  speak  with  confi- 
dence as  to  the  far  greater  variations  of  the  latter. 

To  determine  just  how  many  of  the  North  Pacific  forms  are 
really  "  species,"  or  even  reliable  subspecies  or  varieties,  is  not  pos- 
sible, at  present.  Perhaps  it  never  will  be  possible,  though  much 
nearer  approaches  to  the  solution  of  the  problem  may  be  made  here- 
after, aided  by  more  studies  of  the  forms  in  life,  and  by  far  more 
extensive  collections. 

Professor  Fisher  thinks,  as  I  have  long  believed,  that  many  of 
the  puzzling  forms  are  mere  hybrids  between  a  few  associated 
species.  If  so,  some  of  the  hybrids  may,  by  isolation,  etc.,  have 
formed  more  or  less  fixed  local  varieties,  or  even  real  species,  in 
modern  times,  while  others  may  be,  at  present,  mere  sporadic  or 
individual  cases.  Much  larger  collections,  from  wider  areas,  might 

*Many  years  ago  (op.  cit,  1894),  I  gave  a  full  description  of  H.  pectinata, 
a  supposed  second  species  from  New  England.  Later  I  reduced  it  to  a 
variety  of  H.  sanguinolenta.  Yet  Dr.  Fisher  quotes  it  (IQII&,  p.  10)  as  a 
"nomen  nudum."  The  same  form  occurs  in  Bering  Sea.  See  below,  and  pi. 
XLIX,  figs,  i,  10.  Another  form  (H.  eschrichtii  Miiller  and  Troschel)  was  for- 
merly described  (1842)  from  Greenland,  but  it  has  almost  invariably  been  con 
sidered,  and  as  I  believe  correctly,  a  slight  variation  of  typical  sanguinolenta. 
Dr.  Fisher  has  applied  the  name  to  what  I  consider  entirely  different  forms, 
from  the  North  Pacific. 


SHALLOW-WATER   STARFISHES  213 

well  enable  us  to  distinguish  sporadic  forms  from  more  fixed  races, 
even  among  hybrid  forms. 

At  present,  we  can  only  describe  the  apparently  more  important 
forms,  as  they  occur  in  our  collections.  Where  so  many  forms 
exist,  it  is  not  strange  that  each  collection,  of  any  great  size,  should 
give  diverse  results,  even  could  the  "  personal  equation "  be 
eliminated. 

However,  there  are  certain  leading  forms  in  that  fauna  that  all 
investigators  agree  upon,  as  well  established  species. 

Dr.  Fisher  has  also  described  and  figured  several  very  distinct 
species  from  the  deeper  water,  that  do  not  come  within  the  bounds  of 
this  report,  such  as  H.  polyacantha,  H.  clarki,  H.  aleutica,  H.  asthen- 
actis. 

Aside  from  those,  he  recognizes  (pp.  267,  303)  eight  principal 
species  and  named  varieties  from  the  shallow  waters.  The  large 
collections  that  I  have  studied  make  it  seem  necessary  to  add  to 
those,  four  or  five  additional  forms,  worthy  of  names. 

Of  the  dozen  or  so  of  named  forms,  I  believe  that  about  five  or  six 
represent  genuine  species,  besides  the  four  deep-water  species  men- 
tioned above.  But  most  of  the  shallow-water  forms  from  south  of 
Puget  Sound  may  be  referred  to  H.  leviuscula. 

In  a  few  cases  my  own  determinations  differ  from  those  of  Pro- 
fessor Fisher,  especially  in  respect  to  the  arctic  forms;  but  in  the 
main  I  believe  he  has  treated  the  forms  very  judiciously.  His  col- 
lections from  that  coast  were  larger  than  mine.  His  descriptions 
are  detailed  and  his  illustrations  numerous  and  excellent.  To  dupli- 
cate them  would  be  superfluous.  Therefore  his  work  should  be  con- 
sulted by  everyone  interested  in  the  group,  and  especially  for  the 
doubtful  forms.  The  following  details  are,  therefore,  abbreviated 
in  many  cases. 

It  is  not  improbable  that  some  of  the  notable  variations  are  mere 
individual  variations  of  variable  species,  without  any  constancy. 
Other  variations  may  be  due  to  seasonal  changes  in  nutrition  and 
rapidity  of  growth.  I  am  disposed  to  think  that  the  marked  differ- 
ences frequently  noted  in  the  roughness  or  thorny  character  of  the 
tips  of  the  spinules  of  all  the  species  may  be  due  to  changes  in  the 
rapidity  of  growth.  Those  that  have  the  tips  of  the  spinules  spicu- 
lose  or  thorny  may  be  growing,  as  to  their  spines  at  least,  more 
rapidly  than  those  individuals  with  more  even  or  rounded  tips,  as  in 
those  of  typical  leviuscula  and  var.  lunula.  Such  differences  of 
growth  may  be  due  to  the  abundance  or  scarcity  of  food,  or  to 


214  VERRILL 

seasonal  changes  in  the  temperature  of  the  water,  or  to  the  partial 
suspension  of  growth  during  the  breeding  season,  and  the  lack  of 
nourishment  in  the  case  of  the  female,  while  incubating  the  eggs 
and  young.  Moreover,  I  have  good  reason  to  believe  that  in  some 
species  there  may  be  marked  sexual  differences,  though  my  material 
is  not  sufficient  to  determine  this  with  certainty. 

It  is  known  that  in  many  species  the  female,  while  carrying  the 
eggs  and  young,  under  the  mouth  and  oral  part  of  the  disk,  arches 
up  the  abactinal  side  of  the  disk  to  a  considerable  extent,  and  makes 
a  central  cavity  beneath,  in  which  the  eggs  are  held.  (See  Fisher's 
figure,  pi.  LXVIII,  fig.  i.)  In  some  species  this  seems  to  be  only  a 
temporary  condition,  disappearing  at  other  periods.  But  in  H.  tumida 
V.,  of  Alaska,  it  seems  to  have  become  a  permanent  condition,  in  an 
exaggerated  form,  for  the  oral  region  and  mouth  are  deeply  with- 
drawn and  a  number  of  the  oral  plates  have  become  modified,  to  fit 
this  condition.  Whether  all  such  individuals  are  females  I  do  not 
know.  The  male  may,  perhaps,  be  quite  different  in  form. 

•Variations  in  the  slenderness  of  the  rays  seem  to  be  of  little 
importance  in  this  genus,  unless  of  great  extent.  The  thickness  of 
the  rays  at  base  depends  partly  on  the  number  of  rows  and  extent 
of  rows  of  the  interactinal  and  intermarginal  plates,  but  also  upon 
the  number  and  size  of  the  abactinal  ossicles.  These  are  constantly 
increasing  in  number  by  the  interpolation  of  new  ones,  and  probably 
by  the  splitting  up  of  the  older  and  larger  ones,  while  at  the  same 
time  the  smaller  ones  are  growing  larger.  Where  such  large  nunv 
bers  of  ossicles  exist,  a  slight  change  in  the  size  or  form  of  each  will 
cause  great  changes  in  the  whole  skeletal  structure. 

That  variations  in  growth,  more  or  less  periodically,  may  account 
for  many  apparent  varieties,  has  been  notably  confirmed,  for  me,  by 
finding  certain  specimens  of  H.  sanguin&lenta,  with  rays  equal  in 
size  and  length,  but  having  the  marginal  and  other  ossicles  and 
spinules  as  different,  on  some  of  the  rays  compared  with  others,  as 
they  usually  are  in  different  "  varieties  "  of  the  species,  not  only  in 
size,  but  also  in  the  form  and  number  of  spinules  upon  them.  The 
variations,  as  between  adjacent  rays,  may  be  as  much  as  fifty  per 
cent  in  the  size  of  inferomarginal  plates  and  number  of  their  spinules. 
These  are  not  cases  of  restorations  after  injury,  which  often  also, 
cause  peculiar  abnormal  variations. 

The  habit  of  carrying  the  eggs  and  young  till  well  developed,  as  in 
Leptasterias  and  Solaster,  is  conducive  to  variations  perpetuated  by 
isolation  of  colonies,  due  to  their  slow  rates  of  diffusion.  The 


SHALLOW-WATER   STARFISHES  215 

remarks  on  this  subject  under  Leptasterias  will  no  doubt  apply  just 
as  well  to  Henricia,  for  the  young  are  carried  in  the  same  way. 
(See  pp.  116,  117,  above.) 

Consequently,  it  is  not  at  all  surprising  that  numerous  more  or  less 
localized  varieties  of  Henricia  should  be  found  on  the  vast  extent  of 
coast-line  from  Puget  Sound  to  the  Arctic  Ocean.  The  wonder 
rather  is  that  any  of  the  species  or  varieties  should  preserve  their 
essential  characters,  as  several  of  them  do,  over  such  extensive 
regions  as  the  whole  coast-line  from  Vancouver  Island  to  the  Aleutian 
Islands,  for  example,  or  even  from  California  to  Unalaska. 

HENRICIA  LEVIUSCULA  (Stimpson)  Fisher. 

Plate  xn,  figures  5,  6;  plate  xiu,  figures  i,  2;  plate  LXXXVIII,  figures  i,  10, 

2-2C  (varieties). 

Linckia  leviuscula  STIMPSON,  Journ.  Boston  Soc.  Nat.  Hist,  vi,  p.  529  [p.  89], 

1857- 
Cribrella  leviuscula  VERKILL,  Trans.  Conn.  Acad.,  i,  p.  326,  1867.    Sladeti,  Voy. 

Challenger,  xxx,  pp.  542,  806  (distribution  only).    Ives,  Proc.  Acad.  Nat. 

Sci.  Philadelphia,  1889,  p.  169  (distribution  only).    De  Loriol,  Mem.  Soc. 

Phys.  Nat.  Geneve,  xxxn,  No.  9,  p.  14,  pi.  n,  figs.  2-2c,  1897  (description 

only).    Clark,  Proc.  Boston  Soc.  Nat.  Hist.,  xxix,  p.  327,  1901  (varieties). 
Henricia  leviuscula  FISHER,  1910,  p.  570;  19116,  p.  280,  pi.  LXIX,  figs,  i,  2;  pL 

LXX,  figs.   I,  2J   pi.  LXXI,  figs.  2,  3J  pi.  CXI,  fig.  6. 

Rays  five,  long  and  terete,  gradually  tapered.  Disk  rather  small. 
Radii  of  an  average  dry  specimen,  8  mm.  and  42  mm. ;  ratio,  about 
i :  5.25.  Breadth  of  ray  at  base,  9  mm.  A  larger  specimen  has  the 
radii  12  mm.  and  75  mm. ;  ratio,  i :  6.25.  It  grows  much  larger 
than  this.  One  example  from  Alaska  has  the  radii  15  mm.  and 
85  mm.  Many  specimens  have  more  slender  rays,  others  have  them 
stouter  than  those  given. 

The  dorsal  ossicles  are  thick  and  pretty  closely  united  in  a  fine- 
meshed  reticulation,  leaving  very  small  papular  spaces  between 
them ;  the  papulae  are  mostly  isolated  in  small  specimens,  but  three  to 
five  together  in  large  ones.  The  ossicles  are  convex,  unequal, 
rounded,  elliptical,  curved  lunate,  or  short  cordate.  The  convex 
spinulose  surface  is  often  crescent-shaped.  The  clusters  of  spinules 
are  usually  densely  crowded,  varying  much  in  size  and  form,  as  do 
the  ossicles;  but  the  greater  number  in  the  typical  variety  are 
rounded,  or  short  elliptical,  with  the  longer  axis  transverse  to  the 
rays  or  often  oblique. 

The  spinules  are  very  small,  short,  nearly  even,  usually  with 
minutely  spinulose  tips.  The  two  marginal  and  the  peractinal  rows 


2l6  VERRILL 

of  ossicles  and  spines  are  very  distinct  and  regular,  running  parallel 
nearly  to  the  base  of  the  rays ;  but  the  peractinals  are  rather  smaller 
and  slightly  more  numerous,  so  that  not  all  are  opposite  the  mar- 
ginals. The  ossicles  of  the  two  marginal  rows  are  opposite,  the 
inferior  ones  a  little  larger  and  more  rectangular.  The  spiniferous 
surface  is  broadly  elliptical  or  squarish,  bearing  dense,  broad  ellipti- 
cal or  quadrilateral  clusters  of  spinules,  slightly  larger  than  the 
dorsals,  and  not  much  longer  transversely  than  longitudinally.  The 
clusters  of  spinules  on  the  peractinal  ossicles  are  rather  smaller  and 
more  rounded,  but  distinct  nearly  to  the  tips  of  the  rays,  where  they 
become  very  small.  The  upper  and  lower  marginal  rows  diverge 
rather  abruptly,  close  to  the  bases  of  the  rays,  and  two  or  three  very 
short  rows  of  smaller  ossicles  are  interpolated,  making  a  small, 
sharply  rhombic,  interradial  area.  The  madreporite  is  spinulose. 

The  adambulacral  ossicles  bear  a  double  or  triple  transverse  group 
of  about  twelve  to  twenty,  or  even  more,  unequal  spines,  increasing 
in  size  and  length  from  the  exterior  to  the  margins  of  the  groove. 
The  outer  ones  are  very  small,  not  larger  than  the  peractinals ;  the 
odd  one  on  the  margin  of  the  groove  is  larger  than  any  of  the  others ; 
within  the  groove  there  is  a  small  and  very  short  spine. 

The  adoral  and  oral  spines  are  small,  and  not  unlike  the  other 
adambulacrals,  but  the  perorals  and  epiorals  are  usually  a  trifle 
stouter  than  the  rest.  (See  pi.  xm,  fig.  2.) 

In  life  the  color  is  usually  orange  or  orange-red,  but  varies  to  a 
variety  of  other  colors,  including  dark  red  and  purple.  It  is  often 
mottled  with  red  or  brown  on  a  ground-color  of  yellow,  orange,  drab, 
light  brown,  pink,  or  lavender. 

This  species  is  common  in  shallow  water  and  at  low  tide,  from 
Monterey,  California,  to  Yakutat,  Alaska. 

I  have  examined  specimens  from  San  Luis  Obispo  Bay ;  Monterey ; 
San  Francisco;  Tomales  Bay;  Victoria,  Vancouver  Island;  Queen 
Charlotte  Islands;  Sitka;  and  many  other  places.  It  was  taken  by 
the  Harriman  Expedition  at  Sitka,  Yakutat,  Kadiak,  Orca,  etc. 
Formerly  recorded  from  Puget  Sound  (Stimpson,  Clark,  etc.)  ;  Mar- 
mot Island  (Ives,  1889) ;  Vancouver  Island  (De  Loriol)  ;  California, 
etc.  Mr.  Fisher  (19116)  gives  its  range  as  from  the  Aleutian 
Islands  to  San  Diego,  California,  and  from  low  tide  to  eighty 
fathoms,  but  most  of  his  localities  are  south  of  Sitka,  and  in  less 
than  fifty  fathoms. 


SHALLOW- WATER   STARFISHES  217 

VARIATIONS. 

This  species  varies  considerably  in  the  proportions  of  the  rays 
and  disk,  some  specimens  having  very  long  and  slender  rays,  with  a 
small  disk,  while  others  have  the  rays  shorter  and  more  robust,1 
much  as  in  H.  sanguinolenta.  There  is,  also,  considerable  variation 
in  size  and  form  of  the  dorsal  ossicles,  some  of  the  specimens  almost 
lacking  the  peculiar  curved  forms,  characteristic  of  most.  The 
dorsal  spinules,  always  numerous,  vary  in  size,  so  that  the  degree  to 
which  they  are  crowded  is  variable.  Commonly  they  are  obtuse,  or 
clavate,  but  often  they  are  acute,  or  have  thorny  or  spiculose  tips. 
The  number  and  size  of  the  adambulacral  spines  are  also  variable. 
But  the  species  is  usually  easily  recognized. 

Six-rayed  specimens  occur  rarely  on  the  California  coast. 

Professor  Fisher  (op.  cit,  19116,  p.  283)  mentions  a  dwarf  lit- 
toral variety  found  at  Monterey,  California,  which  carried  clusters  of 
orange-colored  eggs  in  January.  The  disk  was  arched,  making  the 
oral  region  concave,  as  usual  in  this  genus  when  eggs  or  young  are 
carried  by  the  mother. 

The  following  eight  varieties,  from  California  to  Sitka,  are  not 
regarded  as  subspecies  by  me,  but  rather  as  local  varieties. 

HENRICIA  LEVIUSCULA,  Var.  LEVIUSCULA   (Stimpson). 
Plate  xu,  figure  5 ;  plate  xni,  figures  I,  2. 

The  specimen  in  the  U.  S.  National  Museum,  supposed  to  be 
Stimpson's  type,  is  a  small,  slender-rayed  specimen  in  rather  poor 
preservation. 

It  belongs  to  the  common  variety  having  rather  large,  well-rounded 
dorsal  pseudopaxillae,  pretty  close  together,  x  and  with  small  papular 
areas.  The  paxillary  spinules  are  very  short,  even,  and  numerous. 

The  two  rows  of  marginals  and  the  interactinal  row  of  plates  are 
very  distinct  and  pretty  regular. 

The  inferomarginals  are  the  larger  and  are  squarish.  The  adam- 
bulacral spines  are  rather  numerous ;  the  inner  two  to  four  are  the 
larger. 

lProf.  H.  L.  Clark  (op.  cit,  p.  327,  1901)  proposed  varietal  names  based  only 
on  the  relative  length  of  the  rays  and  size  of  the  disk.  The  stouter-  and 
shorter- rayed  form  (radii  about  i :  2-3.5)  he  called  var.  crassa;  the  longer- 
rayed  form  (radii  about  1:5-6),  var.  attenuata.  But  as  all  grades  of  inter- 
mediate forms  occur,  these  variations  cannot  be  regarded  as  of  much 
significance.  Several  other  species  of  the  genus,  if  not  all,  vary  in  the  same 
manner. 


2l8  VERRILL 

I  have  seen  many  specimens  that  agree  well  with  Stimpson's  typical 
form.  They  are  peculiar  mainly  in  having  the  larger  dorsal  ossicles, 
round  or  elliptical,  and  covered  with  compact  clusters  of  stout  clavate 
or  obtuse  spinelets,  so  crowded  together  as  to  give  the  groups  an 
evenly  convex  form.  They  are  peculiar,  also,  in  having  fewer  and 
larger  adambulacral  spines  than  in  some  other  varieties.  These  stand 
in  two  transverse  rows  of  about  six  to  eight  each,  increasing  from  the 
outer  to  the  furrow  end  of  the  plate,  but  even  the  outer  ones  are  as 
large  as  the  adjacent  peractinal  spines,  while  the  marginal  odd  spine 
is  relatively  large  and  stout;  the  single  spine  within  the  groove  is 
small  and  slender. 

Monterey,  San  Francisco,  and  other  localities  on  the  coast  of 
California  and  northward  to  British  Columbia.  The  three  figures 
are  all  from  one  specimen,  from  Monterey. 

This  seems  to  be  a  southern  variety  of  the  species.  The  increased 
thickness  of  the  ossicles  and  spines  may  be  due  to  the  warmer  climate, 
or  to  a  larger  amount  of  lime  in  the  food. 

HENRICIA  LEVIUSCULA,  Var.  LUNULA  Verrill,  nov. 
Plate  LXXXVIII,  figures  2-2<r  (details). 

This  is  one  of  the  most  common  shore  starfishes  from  California  to 
middle  Alaska. 

Rays  five,  variable  in  length  and  thickness,  usually  terete  and 
regularly  tapered. 

The  marginal,  interactinal,  and  adambulacral  plates  and  spinules 
are  essentially  as  in  the  typical  leviuscula;  and  as  in  that  form  the 
marginals  and  interactinals  form  three  very  evident,  regular,  longi- 
tudinal rows  of  imbricated  plates  that  are  larger  and  wider  than  the 
rest,  and  covered  with  large  numbers  of  spinules. 

The  only  notable  feature  for  distinction  is  found  in  the  forms  of 
the  principal  abactinal  pseudopaxillae  and  ossicles.  These,  instead  of 
being  nearly  round  or  broad  elliptical,  as  in  typical  leviuscula,  are 
curved  in  a  more  or  less  crescent  form  with  blunt  cusps,  or  may  be 
said  to  have  an  elongated  reniform  shape,  partially  surrounding  and 
enclosing  the  papular  pore  on  the  concave  side,  which  is  adoral. 

Although  this  peculiarity  appears  to  be  of  minor  importance,  it 
gives  a  special  fades  to  the  variety,  easily  recognized  even  by  a 
superficial  examination. 

Large  numbers  of  this  form  have  been  examined,  from  Vancouver 
Island,  Queen  Charlotte  Islands,  California,  etc.,  and  some  from 
Sitka  and  Dutch  Harbor.  The  specimen  affording  the  figures  of  the 
ossicles  was  from  Monterey,  California. 


SHALLOW-WATER   STARFISHES  219 

This  variety  often  intergrades  with  the  typical  form  and 
with  other  varieties.  It  is,  perhaps,  the  predominant  form  of  the 
species,  in  shallow  water,  when  well  grown,  but  the  original  form, 
described  by  Dr.  Stimpson,  had  roundish  abactinal  ossicles. 

In  many  cases  the  longer,  lunulate  ossicles  show  one  or  more 
lines  or  grooves  across  them,  apparently  indicating  a  process  of 
dividing  into  two  or  more  shorter  parts.  Should  such  a  process  occur 
extensively,  it  would  convert  an  individual  of  this  variety  into  the 
typical  form.  This  may,  perhaps,  occur  periodically,  or  at  the  times 
of  more  rapid  growth. 

HENRICIA  LEVIUSCULA,  Var.  ATTENUATA  (Clark). 

Cribrella  leviusculo,  var.  attenuata  H.  L.  CLARK,  Proc.  Boston,  Soc.  Nat 
Hist.,  xxix,  p.  327,  1901. 

Some  specimens  from  Departure  Bay,  British  Columbia,  are 
remarkable  for  the  slenderness  of  the  rays.  The  most  attenuated 
has  the  radii  7  mm.  and  50  mm. ;  ratio,  1:7.  In  another  they  are 
9  mm.  and  54  mm. ;  ratio,  1 : 6.  In  the  former  the  breadth  of  the 
ray  in  the  middle  is  5  mm.;  in  the  latter  it  is  6  mm.  These  are, 
therefore,  much  more  attenuated  than  the  one  named  by  Dr.  Clark. 

The  rays  are  terete  and  evenly  tapered  from  close  to  the  base. 
The  dorsal  pseudopaxillae  are  roundish,  pretty  uniform  in  size,  even, 
and  crowdedly  covered  with  minute  spinules. 

The  two  marginal  rows  of  plates  are  very  evident,  their  plates 
are  squarish,  about  equal  in  size  to  dorsal  ones,  and  covered  with 
similar  spinules.  One  row  of  actinal  plates,  similar  in  size  proxi- 
mally,  extends  to  the  distal  fourth  of  the  ray;  their  spinules  are  a 
little  longer  and  coarser.  Adambulacral  spines  are  slender,  ten  to 
twelve  to  a  plate ;  the  inner  ones  are  larger. 

The  color  is  orange  and  orange-red  above ;  yellow  below. 

Departure  Bay,  British  Columbia  (C.  H.  Young,  Geological  Sur- 
vey of  Canada)  ;  Puget  Sound  (Clark)  ;  etc. 

HENRICIA  LEVIUSCULA,  var.  INEQUALIS  Verrill,  nov. 
Plate  LXXXVHI,  figures  I,  ia. 

The  type  is  rather  large,  with  five  long,  regularly  tapered,  mod- 
erately slender  rays.  The  radii  are  15  mm.  and  83  mm. ;  ratio,  1 :  5.5. 

The  dorsal  surface  is  closely  covered  with  small  parapaxillae,  mostly 
roundish  or  elliptical,  sometimes  slightly  curved,  and  bearing  numer- 
ous small,  divergent,  often  stellate,  very  slender,  partly  clavate, 


220  VERRILL 

rough-tipped  spinules,  much  like  those  of  var.  dyscrita.  The  papular 
pores  are  numerous  and  rather  small ;  the  areas  are  mostly  narrower 
than  the  plates. 

The  two  rows  of  marginal  plates  and  the  peractinal  row,  on  the 
proximal  part  of  the  ray,  are  regular  and  subequal,  transversely  elon- 
gated, much  like  those  of  var.  leviuscula  in  form  and  size;  close  to 
the  base  of  the  ray  there  may  be  a  few  small  subactinal  plates,  and 
two  short  rows  of  intermarginals.  All  these  plates  are  covered  with 
slender,  rough-tipped,  divergent  spinules  like  those  of  the  dorsal 
side,  but  longer ;  there  may  be  fifty  or  sixty  on  the  larger  marginal 
plates,  of  which  about  twenty-five  are  on  the  border. 

The  adambulacral  spines  are  the  most  characteristic  feature.  They 
are  all  much  larger  and  longer  than  in  the  typical  variety,  mostly 
terete,  slightly  clavate,  obtuse.  There  are  mostly  but  ten  to  twelve, 
paired  in  two  rather  regular,  divergent  rows,  the  inner  one  or  two 
only  a  little  larger.  The  rest  are  graded,  but  the  outermost  are  more 
than  twice  as  large  as  those  of  the  marginals. 

The  type  is  from  the  Queen  Charlotte  Islands  (coll.  G.  M.  Dawson, 
Geological  Survey  of  Canada),  received  in  exchange  from  J.  F. 
Whiteaves  (No.  5133,  Yale  Mus.).  Others,  very  similar,  are  from 
off  Victoria  and  Sitka. 

HENRICIA  LEVIUSCULA  SPICULIFERA  (Clark). 
Plate  LXXXVII,  figures  2-2& ;  plate  cvn,  figure  3. 

Cribrella  spiculifera   CLARK,   Proc.    Boston    Soc.    Nat.   Hist.,   xxix,   p.   328, 

pi.  n,  figs,  i,  2;  pi.  iv,  fig.  i,  1901. 
Henricia  leviuscula  multispina   (pars')  FISHER,  op.  cit.,  1910,  p.  571;   191  ib, 

p.  286,  pis.  LXXII,  LXXIII. 
Henricia  spiculifera  FISHER,  op.  cit.,  19116,  p.  295  (copied  from  Clark). 

The  two  type-specimens  of  this  subspecies  or  variety,  which  were 
in  the  collections  of  Columbia  University,  have  been  examined  by  the 
writer.  They  were  preserved  in  alcohol.  Radii,  in  one  example, 
about  14  mm.  and  64  mm.;  in  the  other,  15  mm.  and  77  mm. 
Ratios,  about  as  1:5.  The  five  rays  are  rather  stout  and  swollen  at 
base,  tapered  distally  to  rather  slender  tips. 

In  these  specimens,  which  were  somewhat  relaxed  or  flaccid  in 
alcohol,  the  dorsal  ossicles  are  not  closely  crowded,  but  seemed 
separated  by  intervening  spaces,  the  visible  portion  being  prominent. 
The  papular  areas  appear  rather  large.  The  dorsal  ossicles  are 
small,  and  very  unequal  in  size  and  form  and  show  no  definite 
arrangement  in  radial  rows.  Many  of  the  larger  ones  are  promi- 
nently crescent-shaped,  as  exposed ;  others  are  oblong  or  elliptical ; 


SHALLOW- WATER   STARFISHES  221 

the  smaller  ones  are  rounded  or  elliptical.  All  are  crowdedly  cov- 
ered with  very  numerous  fine  and  slender,  rather  long,  spicule-like, 
equal  spinules. 

The  upper  and  lower  rows  of  marginal  plates  are  easily  recog- 
nizable, the  lower  ones  larger.  They  are  distinctly  larger  than  the 
adjacent  dorsals,  but  bear  the  same  kind  of  slender  spinules.  They 
are  transversely  oblong-elliptical  in  form,  becoming  more  quad- 
rangular distally,  where  the  two  rows  are  contiguous  and  similar. 
Proximally  they  diverge  strongly  and  are  separated  at  the  base  of 
the  rays  by  two  or  more  short  rows  of  intermarginal  plates,  similar 
in  form  to  the  supramarginals,  but  rather  smaller;  of  these,  the 
middle  row  may  extend  to  about  the  mid-length  of  the  ray. 

A  single  row  of  smaller  interactinal  parapaxillae  (sometimes 
double)  intervenes  between  the  lower  marginals  and  adambulacrals. 
They  are  somewhat  quadrangular  or  roundish  in  form  and  extend 
nearly  to  the  tips  of  the  rays ;  they  bear  fine  spinules,  like  those  of  the 
marginals.  The  papulae  between  the  marginal  and  peractinal  plates 
are  rather  large  and  mostly  stand  isolated. 

The  adambulacral  plates  bear  an  unusually  large  number  of 
spinules,  often  twenty-five  to  thirty  or  more.  These  spinules  are 
slender,  erect,  and  closely  crowded,  decreasing  in  size  and  length 
from  the  margin  of  the  groove  outwardly;  the  outer  ones  are 
similar  in  size  to  those  of  the  adjacent  interactinal  plates.  They 
mostly  stand  in  three  or  four  radial  rows  on  each  plate,  while  there 
may  be  six  to  nine  in  each  transverse  row.  The  two  or  three  standing 
at  the  margin  of  the  groove  are  distinctly  larger  than  the  rest ;  and 
next  to  these  is  a  second  oblique  row,  usually  of  three,  that  are 
smaller,  but  distinctly  larger  than  the  third  row.  The  single  furrow- 
spine  is  short,  acute,  and  hardly  reaches  the  margin.  The  adambu- 
lacral spines  are  all  slender  and  not  clavate. 

Adoral  and  epioral  spines  numerous,  crowded,  scarcely  larger 
than  adjacent  adambulacrals,  and  similar  in  form,  seven  to  nine  in 
each  marginal  series. 

The  general  appearance  of  this  variety  is  similar  to  that  of  a 
stout-rayed  H.  leviuscula  or  sanguinolenta,  but  it  is  peculiar  in  hav- 
ing unusually  large  numbers  of  crowded  and  slender  adambulacral 
spines  on  each  plate.1  A  larger  series  of  specimens  shows  apparently 
intermediate  forms  between  this  and  leviuscula. 

*Dr.  Clark's  diagrammatic  figure  (pi.  iv,  fig.  i)  represents  them  as  much 
more  regular  and  equal  than  they  actually  are.  The  exterior  ones  are  rep- 
resented as  about  as  long  as  the  inner  or  marginal  ones,  but  they  are  not 
half  as  long. 


222  VERRILL 

After  a  careful  study  of  the  type  I  cannot  see  any  good  reasons  for 
considering  spiculifera  a  real  species,  distinct  from  leviuscula. 
Some  of  the  differences  are  due  to  the  somewhat  flaccid  condition  of 
the  alcoholic  specimens.  Intermediate  specimens  are  of  frequent 
occurrence  in  the  same  region. 

Puget  Sound  (Expedition  of  Columbia  University,  1896-7). 

Other  good  examples  are  from  Kadiak,  5  fathoms,  and  Sitka 
( W.  R.  Coe,  Harriman  Expedition) . 

A  specimen  similar  to  the  above,  in  many  respects,  has  been  sent 
to  me  by  the  U.  S.  National  Museum.  It  was  taken  at  Bering 
Island  by  Mr.  N.  Grebnitsky.  It  is  the  specimen  figured  (pi.  cvn, 
fig.  3;  and  pi.  LXXXVII,  figs.  2,  20). 

I  believe  that  many  of  the  intermediates  are  the  same  as  spiculi- 
fera, which  has  a  decided  resemblance  to  leviuscula  dorsally.  The 
more  arctic  specimens  may  have  had  a  different  origin,  and  belong 
with  a  different  species,  more  nearly  related  to  sanguinolenta.  The 
four  specimens  figured  by  Fisher  are  all  from  the  Aleutian  Islands 
and  Kadiak,  within  the  known  range  of  leviuscula. 

I  should  refer  most  of  his  figured  specimens  to  spiculifera,  judging 
from  the  description  and  figures,  which  are  very  good.  Perhaps 
it  is  merely  the  normal  cold  water  form  of  leviuscula. 

HENRICIA  LEVIUSCULA,  Var.  MULTISPINA  Fisher. 

Henricia  multispina  FISHER,  op.  cit.,  1910,  p.  571 ;  191  ib,  p.  286  (pars)  ;  pis. 
LXXII  and  LXXIII  (pars). 

H.  multispina  Fisher,  as  figured,  differs  but  slightly,  and  perhaps 
ought  to  be  called  the  same  as  spiculifera.  Some  of  the  differences 
given  by  Fisher,  on  his  p.  297,  do  not  hold  good,  being  partly  due 
to  the  different  states  of  preservation.  This  applies  particularly  to 
the  dorsal  skeleton.  The  arrangement  of  the  adambulacral  spines  is 
essentially  the  same,  but  Clark's  diagrammatic  figure  is  misleading. 
Such  differences  as  actually  exist  seem  trivial  and  such  as  are  notori- 
ously variable. 

The  particular  point  which  makes  the  full  identity  doubtful  is  the 
fact  that  some  of  Dr.  Fisher's  specimens  of  multispina  came  from 
Bering  Sea  and  Siberia,  far  north  of  the  ordinary  range  of  leviuscula. 
The  type  was  from  the  Aleutian  Islands.  Fisher  himself  states,  on 
pp.  288,  289,  that  large  numbers  of  his  specimens  from  the  shore 
and  very  shallow  water,  south  of  the  Aleutian  Islands,  at  Bristol 
Bay,  Kadiak,  etc.,  are  intermediate  between  the  type  and  leviuscula. 
However,  Dr.  Fisher  himself  records  leviuscula  from  Bering  Sea. 


SHALLOW-WATER   STARFISHES  223 

I  have  seen  no  specimens  of  it  from  much  north  of  the  Aleutian 
Islands,  which  are  usually  considered  its  northern  limit. 

I  have  studied  specimens  from  Bering  Sea,  which  are,  perhaps,  of 
the  same  variety  as  some  of  those  recorded  by  Fisher  from  the  same 
region.  My  arctic  form,  however,  appears  to  be  a  variety  of  H.  san- 
guinolenta,  which  is  much  like  .the  Atlantic  variety  pectinata. 

Professor  Fisher  gives  the  range  of  his  multispina  as  from  Oregon 
to  Bering  Strait  and  Kuril  Islands,  and  from  low  tide  to  238  fathoms. 

This  species  or  subspecies  appears  to  be  closely  related  to,  and  may 
be  the  same  as  Henricia  densispina  (Sladen,  1878,  p.  432,  pi.  vm, 
figs.  5-9,  as  Cribrella),  from  the  Straits  of  Korea,  40  fathoms. 

The  latter  also  has  two  spines  on  the  furrow  margin  and  several 
graded  pairs  back  of  them,  with  very  numerous,  crowded,  rough, 
thorny  spinules  on  the  close  marginal  and  dorsal  plates ;  papular  areas 
small,  mostly  with  isolated  or  few  papulae. 

The  Japanese  species,  figured,  but  not  described,  by  Ives  (op.  cit., 
1891,  p.  212,  pi.  ix,  figs.  1-4)  as  Cribrella  sanguinolenta,  is  not  that 
species.  It  belongs  to  the  section  of  the  genus  having  a  pair  of  spines 
on  the  furrow  margin,  but  it  has  fewer  on  the  actinal  surface.  Its 
dorsal  ossicles  are  narrow,  very  openly  reticulated,  bearing  spinules 
in  about  two  rows,  and  surrounding  unusually  wide  papular  areas.  It 
may  be  called  HENRICIA  JAPONICA. 

HENRICIA  LEVIUSCULA,  var.  DYSCRITA  Fisher 

(as  a  subspecies). 

Plate  xn,  figure  6. 

Henricia  leviuscula  dyscrita  FISHER,  op.  cit.,  191  ib,  p.  289,  pi.  LXXIV,  figs. 

1-5- 

Form  and  general  appearance  intermediate  between  leviuscula  and 
multispina;  abactinal  ossicles  smaller  than  In  the  latter,  with  similar, 
but  fewer,  delicate  spinules  and  usually  larger  papular  areas,  larger 
than  the  pseudopaxillae.  The  abactinal  spinules  end  in  several  deli- 
cate points. 

Adambulacral  plates  carry  about  fifteen  spinules;  two  intra-fur- 
row  spinules  occur,  if  at  all,  only  near  the  tip  of  the  ray. 

Dr.  Fisher  records  this  form  from  eighteen  localities  in  21  to  80 
fathoms,  from  Monterey  to  Southern  California.  South  of  Monterey 
"  it  predominates  everywhere  over  leviuscula." 

Two  specimens  from  Sitka  (coll.  Harriman  Expedition)  seem  to 
agree  closely  with  this  form  in  all  respects.  (See  pi.  xn,  fig.  6.) 
Others  are  from  British  Columbia. 


224  VERRILL 

HENRICIA  LEVIUSCULA,  Var.  ANNECTENS  Fisher. 

Henricia  leviuscula  annectens  FISHER,  1910,  p.  572;  191  ib,  p.  291,  pi.  LXX, 
fig.  2d;  pi.  LXXI,  figs.  1-3. 

Intermediate  between  leviuscula  and  aspera,  but  superficially 
more  like  the  latter.  Abactinal  skeleton  less  open  than  in  the  latter, 
with  smaller,  deep,  papular  areas.  Ridges  between  papulae  divided 
into  distinct  pseudopaxillas,  the  larger  with  five  to  twenty  short, 
stubby  spinules.  Marginal  plates  more  compressed  than  in  levius- 
cula, with  comparatively  few  spinules.  No  intermarginals  beyond 
first  two  or  three  marginals.  Interactinals  extend  to  one-half  or 
two-thirds  the  length  of  the  ray.  Adambulacral  plates  bear  ten  to 
twelve  spinules  in  two  transverse  rows  and  one  furrow-spine  deep  in 
the  groove,  except  near  the  tip  of  the  ray,  where  there  are  two. 

Dr.  Fisher  records  this  form  from  thirteen  localities  in  21  to  73 
fathoms,  from  Washington  and  Oregon  to  Santa  Barbara,  Cali- 
fornia. 

Specimens  agreeing  very  closely  with  it  were  in  my  collections 
from  California.  The  above  diagnosis  is  condensed  from  that  of 
Dr.  Fisher. 


HENRICIA  LEVIUSCULA,  Var.  SPATULIFERA  Verrill. 

Plate  v,  figure  i;  plate  xiv;  text-figure  12  (details). 
Henricia  spatulifera  VERRILL,  Amer.  Naturalist,  XLIII,  pp.  554,  555,  fig.  6,  1909. 

The  type  of  this  peculiar  variety  is  similar  to  an  average  well- 
grown  robust  specimen  of  leviuscula,  var.  lunula,  in  size  and  form, 
Radii,  13  mm.  and  78  mm. ;  ratio,  1 : 6. 

The  chief  peculiarity  is  in  the  armature  of  the  adambulacral 
plates.  These  bear  a  crowded  group  of  about  eight  to  twelve  spin- 
ules, in  two  close  rows,  which  are  remarkably  large  and  flat,  expand- 
ing distally  to  broad  truncate  tips,  which  are  often  gouge-shaped  or 
grooved  on  one  side.  They  are  so  large  and  crowded  that  they  over- 
lap each  other  and  the  adjacent  plates,  making  it  difficult  to  count 
their  number. 

The  adoral  and  oral  plates  have  the  same  sort  of  armature.  This 
kind  of  spinules  extends  to  the  tips  of  the  rays,  the  size  gradually 
diminishing  to  minute  spatulate  forms  near  the  tips,  with  the 
exception  of  a  limited  section  in  the  middle  of  one  ray,  where  the 
large,  flat  spinules  cease  abruptly  and  the  following  twenty-five 
pairs  of  adambulacral  plates  all  bear  very  small,  short,  slender 
spinules  in  a  crowded  group  of  about  fifty  or  more,  in  about  four 


SHALLOW-WATER   STARFISHES  225 

rows,  twelve  to  fourteen  each,  all  of  nearly  the  same  size.  They 
form  clusters,  as  numerous  or  more  so,  and  of  as  small  size,  as  those 
in  var.  spiculifera  or  multispina,  and  more  uniform  in  size.  After 
these  twenty-five  pairs  of  plates,  those  more  distal  suddenly  resume 
the  same  spatulate  form  and  arrangement  as  on  the  other  rays. 

Probably  these  peculiar  twenty-five  plates  had  been  injured,  losing 
their  spinules,  and  replacing  them  with  the  minute  kind,  just  as  some 
starfishes  will  often  replace  a  lost  spine  of  large  size,  by  a  cluster  of 
granules. 

The  fact  that  a  large  number  of  uniform  small,  slender  spinules 
may  replace  a  group  of  few  large  spatulate  ones,  on  the  same  ray,  is, 
however,  an  important  fact,  showing  that  the  character  of  the  adam- 
bulacral  spinules  is  not  a  matter  of  vital  importance,  and  that  it  might 
easily  vary  more  than  it  usually  does,  even  in  this  variable  genus. 


,m 

a  6       m"1 


FIG.  12. 

Henricia  leviuscula.  var.  spatulifera  Verrill,  type.  Portion  of  under  side;  ad.  adambu- 
lacral  spines;  p,  peractinals;  m,  m",  inferomarginals;  m'".  superomarginals;  ab,  medio-lateral 
pines. 

The  dorsal  skeleton  is  composed  largely  of  elliptical  and  some- 
what lunulate  thickened  ossicles,  rather  loosely  arranged,  leaving 
large  papular  pores,  so  that  the  surface  appears  rougher  than  in  the 
more  typical  varieties.  The  dorsal  spinules  are  small,  mostly  clavate, 
with  thorny  ends,  much  as  in  var.  multispina. 

The  rows  of  marginal  and  peractinal  plates  are  not  so  regular  nor 
so  conspicuous  as  in  most  varieties.  The  inferomarginals  are  the 
largest,  but  not  so  large  as  usual.  The  peractinals  are  much  smaller 
and  roundish. 

The  superomarginals  are  small,  but  larger  than  the  dorso-lateral 
ones,  and  similar  to  the  inferomarginals  in  form.  There  are  two 
rows  of  intermarginals  proximally,  one  being  quite  short.  The 
spinules  on  all  these  plates  are  numerous  and  very  small,  much  as  on 
the  dorsal  side. 
16 


226  VERRILL 

Papular  pores  exist  between  all  these  rows,  and  proximally,  also, 
between  the  peractinal  and  adambulacral  rows. 

The  type  is  from  Monterey,  California  (coll.  Robert  E.  C. 
Stearns,  No.  2238,  Yale  University  Museum).  I  have  examined 
other  specimens  from  Sitka,  Alaska  (Harriman  Expedition). 

Dr.  Fisher  records  a  specimen  from  off  Santa  Cruz  Island,  Cali- 
fornia, in  30  to  41  fathoms  (sta.  4431),  and  two  specimens  from 
Monterey  Bay  shore,  that  may  be  identical  with  this  variety.  He 
puts  them  under  "  variety  E  "  as  "  freaks." 

HENRICIA  SANGUINOLENTA  (Miiller)  Bell. 

Plate  XLIX,  figures  i,  10  (var.  pectinata),  and  figure  2;  plate  LXXXVIII,  figures 
3,  30,  4,  40  (varieties). 

Asterias  sanguinolenta  and  A.  pertusa  MULLER,  Zool.  Dan.  Prod.,  2836  and 

2839,  PP-  234,  235,  1776. 

Asterias  oculata  PENNANT,  Brit  Zool.,  rv,  p.  61,  pi.  xxx,  fig.  56,  1777. 
Asterias  spongiosa  FABRICIUS,  Fauna  Groenl.,  p.  368,  1780.     Gmelin,  p.  3161, 

1788.    Desor,  Proc.  Boston  Soc.  Nat.  Hist.,  in,  p.  67,  1848. 
Cribella  rosea  and  C.  oculata  STIMPSON,  Proc.  Boston  Soc.  Nat.  Hist.,  iv, 

p.  08,  1851. 
Linkia  oculata  FORBES,  Mem.  Wern.  Soc.,  VIH,  p.  120,  pi.  in,  fig.  5,  1839. 

Stimpson,  Invert,  of  Grand  Manan,  p.  14,  1853. 
Henricia  oculata  GRAY,  Ann.  and  Mag.  Nat.  Hist.,  vi,  p.  184,  November,  1840; 

Synopsis,  p.  5,  1866. 

Linkia  pertusa  STIMPSON,  op.  cit,  p.  14,  1853. 
Cribella  oculata  FORBES,  British  Starfishes,  p.  100  (figure),  1841. 
Cribrella  oculata  PERRIER,  Stellerides,  in  Arch.  Zool.  Exper.,  iv,  p.  373,  1875. 

Viguier,  Arch.  Zool.  Exper.,  vn,  p.  128,  pi.  vii,  figs.  8-15,  1878.    Ludwig, 

Echinod.  Mittelm.,  Mittheil.,  Zool.  Station  Neapel,  i,  p.  545,  1879.    Duncan 

and  Sladen,  op.  cit.,  p.  32,  pi.  n,  figs.  18-21,  1881.    Ludwig,  Zool.  Jahrb., 

p.  289,  1886.    Danielssen  and  Koren,  op.  «it,  p.  34,  1884.    Sladen,  op.  cit, 

p.  542,  1889. 
Echinaster  oculatus  MULLER  and  TROSCHEL,  Syst.  Asterid.,  pp.  24,  127,  1842. 

Diiben  and  Koren,  Kongl.  Vetensk.  Akad.  Handlingar,  p.  241,  1844. 
Echinaster  sarsii  MULLER  and  TROSCHEL,  Wiegmann's  Arch.  Nat.,  x,  p.  179, 

1844. 

Echinaster  eschrichtii  MULLER  and  TROSCHEL,  op.  cit.,  1842,  p.  25. 
Echinaster  sanguinolentus  M.  SARS,  Wiegmann's  Arch.  Nat,  x,  p.  169,  1844 

(development).     Fauna  Litt   Norveg.,  I,  p.  47,  pi.  vin,  figs.  3-6,  1846; 

Oversigt  af  Norges  Echinodermer,  p.  84,  1861. 
Cribrella  sanguinolenta  LUTKEN,  Gronl.  Echinod.,  p.  31,  1857.     Verrill,  Proc. 

Boston  Soc.  Nat  Hist.,  x,  p.  345,  1866.   Dujardin  and  Hupe,  Hist.  Nat. 

Zooph.  Echinodermes,  p.  349,  1862.    Norman,  Ann.  and  Mag.  Nat.  Hist., 

Ser.  3,  xv,  p.  24,  1865.  Verrill,  Report  Invert.  Anim.  of  Vineyard  Sd.,  etc., 

in  ist  Annual  Rep.  U.  S.  Fish  Com.,  pp.  407,  420,  447,  496,  719,  1873  (auth. 

ed.,  pp.  113,  425,  etc.,  1874).     A.  Agassiz,  N.  Amer.  Starfishes,  p.   113, 


SHALLOW-WATER  STARFISHES  227 

pi.  xviii,  1877.  Bush,  Proc.  U.  S.  Nat.  Mus.,  p.  246,  1883.  Murdoch, 
Report  Intern.  Boundary  Exped.,  p.  159,  1885.  Ganong,  Echinod.  New. 
Brunswick,  Bull.  Nat.  Hist.  Soc.  New  Bruns.,  vin,  p.  35,  pi  i,  fig.  9, 
1888.  Pfeffer,  Jahr.  Hamb.  Wiss.  Ans.,  1889,  pp.  69,  88,  95,  1890. 
Fewkes,  Proc.  Essex  Inst.,  p.  62,  1891.  ?  Ives,  Proc.  Acad.  Nat.  Sci. 
Philadelphia,  p.  212,  pi.  ix,  figs.  1-4,  1891  (Japan).  Verrill,  Amer. 
Journ.  Sci.,  XLIX,  p.  205,  1895  (distribution).  Ludwig,  Fauna  Arctica, 
I,  p.  472,  1900  (synonymy,  etc.).  Brunchorst,  Bergens  Mus.  Aarbog,  i, 
p.  28,  1902.  Masterman,  Trans.  Royal  Society  of  Edinburgh,  xi,  pp.  373- 
418,  pis.  i-v,  1902  (Abstract  in  Nature,  LXV,  p.  551)  (development). 
Mortensen,  Echinod.  from  E.  Greenland,  p.  72,  pi.  n,  figs.  7-9,  1903 
(varieties).  Clark,  Echinoderms  of  Woods  Hole  Region,  op.  cit.,  1904, 
P.  555.  pl-  HI,  figs.  10,  ii ;  pi.  iv,  fig.  22. 

Echinaster  scrobiculata  DANIELSSEN  and  KOREN,  Norske  Nordhavsexped., 
Nyt.  Mag.  Naturvidsk.,  xxvin,  p.  3,  pis.  i,  n,  1883  (young)  ;  Norwegian 
North  Atlantic  Expedition,  Zool.,  Asteroidea,  p.  40,  pi.  vi,  figs.  10,  n; 
pi.  vn,  figs.  12-14,  1884.  Ludwig,  Fauna  Arctica,  i,  476,  1900  (young). 

Henricia  sanguinolenta  BELL,  Catal.  Echinod.,  p.  95,  1892.  Grieg,  Bergens 
Mus.  Aarbog,  pp.  8,  12,  1896.  Scott,  Proc.  Royal  Phil.  Soc.  Edinburgh, 
p.  189,  1897.  Fisher,  19116,  p.  271,  pi.  LXV,  figs,  i,  2;  pi.  LXVI,  figs.  1-5; 
pi.  LXVIII,  fig.  3.  Coe,  Echinoderms  of  Connecticut,  p.  65,  pi.  i,  figs.  1-3; 
pi  xi,  figs.  1-4,  1912. 

Disk  of  moderate  size  or  small.  Rays  five,  evenly  rounded  in  life, 
rather  long  and  slender,  varying  to  stouter  forms  of  moderate 
length,  usually  evenly  tapered  to  rather  slender  tips. 

The  radii  of  a  well  grown  typical  New  England  specimen  of 
average  proportions,  are  14  mm.  and  62  mm. ;  ratio,  about  i :  4.5 ; 
in  another,  13  mm.  and  58  mm. ;  ratio,  about  i :  4.5.  Many  have 
shorter  rays.  Ambulacral  grooves  are  narrow  and  deep,  often  nearly 
closed  up,  so  that  their  marginal  spines  interlock. 

The  dorsal  and  lateral  skeleton  is  composed  of  a  great  number  of 
small,  nearly  equal,  mostly  rounded  and  elliptical  ossicles,  united 
endwise  into  a  fine-meshed  reticulated  skeleton,  showing  no  median 
row  of  larger  ossicles. 

All  the  ossicles  are  covered  with  divergent  clusters  of  small, 
slender,  rough-tipped  spinules,  nearly  uniform  over  the  dorsal  sur- 
face and  sides  of  the  rays,  and  on  the  disk.  The  papular  areas,  in 
the  meshes,  are  small  and  usually  bear  but  one  to  three  papulae. 

The  madreporic  plate  is  small,  with  only  a  few  rough  gyri,  and 
sometimes  spinulose.  The  pseudo-anal  pore  is  usually  very  distinct 
and  guarded  by  small  incurved  spinules. 

The  superomarginal  and  inferomarginal  ossicles  are  similar  in 
size,  and  form  two  distinct  rows  low  down  toward  the  ventral  side 
of  the  rays.  Distally  they  are  near  together  and  parallel,  closely 
united,  leaving  only  a  simple  row  of  papular  pores  between  them; 


228  VERRILL 

but  on  the  proximal  part  of  the  rays  they  gradually  diverge  and  two 
or  three  short  rows  of  intermarginal  ossicles  are  interpolated  in  the 
interradial  angles. 

The  marginal  plates  in  both  rows  are  a  little  convex  and  lobate. 
The  outer  surface  is  oblong-elliptical,  or  squarish  with  rounded 
corners,  united  by  apophyses.  Their  transverse  diameter  is  often 
but  little  greater  than  the  longitudinal  one,  distally,  but  they  become 
more  transversely  elongated  proximally.  Their  clusters  of  spines 
do  not  much  exceed  those  of  the  dorsal  and  lateral  ossicles,  and 
therefore  they  are  not  so  conspicuous  as  in  H.  leviuscula  and  some 
other  species.  It  often  requires  close  examination  to  distinguish 
them  from  the  adjacent  clusters,  but  in  other  cases  they  are  larger 
and  more  regular,  and  quite  noticeable. 

The  peractinal  ossicles  form  a  regular  row  of  small,  rounded 
plates,  one  standing  opposite  each  adambulacral  and  closely  united 
to  it  and  to  the  opposite  inferomarginal.  This  row  often  disappears 
at  about  the  middle  of  the  ray,  but  may  extend  farther,  sometimes 
nearly  to  the  tip.  The  plates  bear  divergent  clusters  of  spinules, 
usually  smaller  than  those  on  the  adjacent  plates,  but  otherwise 
similar. 

The  adambulacral  plates  mostly  bear  two  transverse  divergent 
series  of  small,  unequal  spines,  usually  standing  in  three  or  four  or 
more  pairs  on  the  external  surface  of  the  plate,  and  regularly 
decreasing  in  size  from  the  inner  toward  the  external  ones.  The 
inner  or  ambulacral  end  of  the  plate  bears  two,  rarely  three,  unpaired 
spines  in  a  median  row.  The  inner  one,  or  furrow-spine,  is  attached 
deep  within  the  furrow  and  usually  projects  horizontally.  It  is  very 
slender  and  acute.  The  outer  one  is  much  larger  and  about  twice  as 
long,  scarcely  tapered,  often  compressed,  blunt  at  the  tip,  and  some- 
times bifid.  It  is  rather  larger  than  the  two  spines  of  the  next 
adjacent  pair.  In  some  large  specimens  many  of  the  larger  adambu- 
lacral spines  are  slightly  capitate  or  clavate.  Distally  they  often 
stand  nearly  all  in  one  median  row  on  each  plate,  but  this  does  not 
often  occur  on  the  proximal  plates. 

Color,  in  life,  variable,  most  frequently  bright  orange-yellow  or 
orange-red,  sometimes  pale  lemon-yellow,  not  rarely  purple. 

Mr.  E.  Desor  (op.  cit,  in,  1848,  p.  n)  not  only  states  that  this 
species  has  the  habit  of  carrying  the  eggs,  but  adds  that :  "  On 
removing  the  eggs  from  the  mother's  embrace,  she  was  seen  to 
move  at  once  directly  towards  and  clasp  them  again." 


SHALLOW-WATER   STARFISHES  229 

The  structure  of  the  skeleton  of  this  species  has  been  beautifully 
illustrated  by  Mr.  A.  Agassiz  (N.  Amer.  Starfishes,  pi.  xvin1),  and 
in  some  particulars  by  Viguier  (1878,  pi.  vn). 

The  distribution  of  H.  sanguinolenta  is  circumpolar.  It  is  known 
from  Greenland  and  the  Arctic  Ocean  generally;  on  the  coasts  of 
northern  Europe  to  Great  Britain,  Ireland,  France,  Biscay  Bay, 
Spitzbergen,  Iceland,  White  Sea,  Barents  Sea,  Kara  Sea,  Okhotsk 
Sea,  etc. ;  also  off  the  Azores.  On  the  northeastern  American  coast 
it  is  very  common  from  Labrador  to  Long  Island  Sound,  in  shallow 
water  (o  to  60  fathoms).  Common  in  the  cold  area  south  of 
Martha's  Vineyard,  in  10  to  60  fathoms.  It  is  also  found  off  Cape 
Hatteras  and  North  Carolina  in  similar  depths.  Taken  at  more  than 
four  hundred  stations  between  N.  lat.  47°  29'  and  35°  38',  by  the 
U.  S.  Fish  Commission.  It  occurs  sparingly  in  the  eastern  part  of 
Long  Island  Sound,  at  Fisher's  Island,  Gardiner's  Island,  and  west- 
ward to  Outer  Island,  near  New  Haven.  Bathymetrical  normal 
range  is  o  to  500  fathoms.  In  the  Bay  of  Fundy  and  on  the  coast 
of  Maine  it  is  common  between  tides.  Off  New  Jersey,  in  1350 
fathoms  (teste  Sladen).  Rare  below  300  fathoms  off  the  Atlantic 
coast.  Faroe  Channel,  125  to  555  fathoms  (Sladen). 

On  the  northwest  coast  of  America  the  typical  form  and  varieties 
have  been  recorded  from  several  localities  in  Bering  Sea  and  Bering 
Strait  and  in  the  adjacent  parts  of  the  Arctic  Ocean,  as  well  as 
from  farther  south.  It  is  probable,  however,  that  many  of  the  more 
southern  shallow-water  records  refer  to  one  of  the  other  species,  or 
to  varieties  of  H.  leviuscula.  I  have  not  seen  it  in  the  collections 
from  southeastern  Alaska,  and  farther  south,  in  shallow  water. 

I  have  examined  Pacific  specimens  from  the  following  places :  Off 
Point  Franklin,  ten  miles,  in  13.5  fathoms,  sand,  1883  (Murdoch, 
U.  S.  Nat.  Mus.,  No.  76231,  variety,  very  large,  dry)  ;  Popoff 
Strait  (W.  H.  Ball,  1872,  one  dry,  var.  pectinata,  No.  561  (924), 
U.  S.  Nat.  Mus.)  ;  Bering  Island  (N.  Grebnitsky,  November,  1889, 
U.  S.  Nat.  Mus.). 

Dr.  Fisher  (19116)  records  it  from  thirty-seven  stations  in  30  to 
344  fathoms,  from  Bering  Sea  to  the  Kuril  Islands;  and  on  the 
American  side  south  to  Washington  (67  fathoms,  one  specimen). 
Nearly  all  of  his  localities  are  in  the  vicinity  of  the  Aleutian  Islands 
or  farther  north. 

Mr.  Ives  recorded  it  from  Japan,  but  his  figures  appear  to  represent 
a  distinct  species.  It  differs  in  its  much  looser  dorsal  reticulations 

1  The  marginal  ossicles  in  his  figs,  i  and  4  are  represented  as  more  regular 
and  symmetrical  than  they  usually  are  in  nature. 


230  VERRILL 

and  smaller  ossicles,  which  bear  small  groups  of  four  to  seven  short, 
blunt  spinules ;  in  the  smaller  size  of  the  inferomarginal  plates,  and 
their  longer  and  larger  spinules;  and  especially  in  the  fewer  and 
unusually  large  adambulacral  spines,  with  a  pair  of  stout  spines 
standing  on  the  margin  of  the  groove,  on  each  plate. 

TERATOLOGY. 

On  the  New  England  coast  I  have  collected  several  specimens 
with  six  regular  rays.  One  of  these,  from  off  Salem,  Massachusetts, 
in  shallow  water,  has  the  rays  uncommonly  slender  and  a  very  small 
disk.  Radii,  7  mm.  and  31  mm. ;  ratio,  about  1 : 4.5.  The  dorsal 
pseudopaxillae  are  very  small  and  bear  mostly  only  two  to  four  spin- 
ules which  are  slender,  thorny  and  acute;  many  have  only  a  single 
spinule.  The  ossicles  are  small,  slender,  and  form  a  fine-meshed 
network. 

The  adambulacral  spines  are  elongated  and  slender,  acute,  about 
four  or  five  in  a  nearly  simple  row. 

The  marginals  of  both  rows  are  small  and  scarcely  different  from 
the  adjacent  laterals  and  peractinals,  but  they  form  pretty  regular 
rows. 

Another  specimen  (Yale  Mus.,  No.  5380),  from  Eastport,  Maine 
(coll.  A.  E.  Verrill,  1870),  is  larger  and  of  about  the  usual  propor- 
tions; radii,  13  mm.  and  58  mm.;  ratio,  about  1 :4-5.  The  spinula- 
tion  is  much  finer  than  that  of  the  typical  form.  The  dorsal  ossicles 
are  much  smaller,  and  roundish,  with  small  papular  areas,  approach- 
ing those  of  var.  miliaris. 

A  specimen  of  moderate  size,  from  Alaska,  has  two  madreporic 
plates,  near  together,  both  interradial;  one  is  more  coarsely  spinu- 
lated;  they  are  about  equal  in  size.  This  specimen  is  rather  more 
coarsely  spinulated  than  usual,  dorsally,  and  the  adambulacral  spines 
are  larger,  finer,  and  less  unequal  than  usual.  Both  rows  of  marginal 
spines  are  unusually  small,  not  larger  than  the  dorso-lateral  ones,  and 
scarcely  distinguishable;  they  have  few  spinules. 

HENRICIA  SANGUINOLENTA,  Var.  PECTINATTA  Verrill. 
Plate  XLIX,  figures  I,  10  (type). 

Cribrella  pectinata  VERRILL,  Proc.  U.  S.  Nat  Mus.,  xvn,  p.  278,  1894. 
Henricia  sanguinolento  variety  C,  FISHER,  op.  cit.,  ignb,  p.  272,  pi.  LXV,  fig.  2; 
pi.  LXVI,  figs,  i,  3,  4,  5. 

Rays  five,  elongated,  rounded,  thick  at  base,  tapering  evenly  to  the 
small  tips.  Disk  moderately  swollen,  the  lesser  to  the  greater  radii 


SHALLOW- WATER   STARFISHES  23! 

of  the  type  as  1 : 4.4.  The  lesser  radius  of  the  type  is  15  mm.;  the 
greater  radius,  66  mm. ;  breadth  of  rays  at  base,  18  mm. ;  diameter 
of  madreporic  plate,  3  mm. 

The  whole  dorsal  surface  and  sides  of  the  rays  are  evenly  covered 
with  small  well  spaced  pseudopaxillae,  each  of  which  bears  a  fascicle, 
or  more  rarely  a  comb-shaped  group  of  four  to  eight  or  more  small 
slender  spinules,  which  stand  nearly  erect,  and  are  nearly  equal  in 
length.  The  pseudopaxillae  arise  from  elevations  of  the  plates  and 
are  so  spaced  as  to  leave  intervals  greater  than  their  own  diameters, 
thus  giving  the  surface  a  rough  papillose  appearance;  the  pseudo- 
paxillae are  more  closely  arranged  on  the  center  of  the  disk  than  on 
the  rays.  The  madreporic  plate  is  large  and  covered  with  rough 
spinules  in  comb-like  groups. 

Each  of  the  interspaces  on  the  rays  bears  a  single  large  papula, 
equal  in  diameter  to  or  exceeding  the  pseudopaxillae ;  similar  papulae 
occur  between  the  ventral  plates,  where  they  form  regular  longitu- 
dinal rows.  On  the  actinal  surface  of  the  rays  there  are  three  regu- 
lar longitudinal  series  of  plates  corresponding  in  number  to  the 
adambulacral  plates.  The  plates  in  the  two  outer  or  marginal  rows 
are  oblong  at  the  summit,  and  each  bears  an  oblong  group  of  slender 
paxilliform  spinules,  arranged  in  two  rows,  and  similar  to  those  of 
the  back.  The  plates  of  the  superomarginal  row  are  somewhat 
smaller  than  those  of  the  inferomarginal,  and  the  spinules  are  about 
twelve  to  fifteen  in  number  toward  the  base  of  the  rays,  while  in 
the  latter  there  are  from  twenty  to  twenty-five  spinules,  which  form 
pretty  regular  comb-like  groups ;  these  extend  to  the  tips  of  the  rays. 
Each  of  the  interspaces  alternating  between  these  rows  of  plates 
contains  a  single  large  papula.  Closely  adjacent  to  the  adambulacral 
plates  there  is  a  row  of  smaller  peractinal  plates,  each  of  which  bears 
a  round  group  of  small  paxilliform  spinules,  ten  to  fifteen  in  number, 
similar  in  size  and  form  to  those  of  the  marginal  plates.  This  row 
of  peractinal  plates  extends  from  the  angle  of  the  jaw  nearly  to  the 
tip  of  the  ray. 

Each  of  the  adambulacral  plates  bears  a  single  small  spine,  situated 
deep  within  and  directly  across  the  furrow,  forming  a  single  longi- 
tudinal series,  and  also  a  transverse  group,  consisting  of  eight  to 
twelve  round,  blunt  spinules,  in  two  rows ;  the  three  inner  ones  are 
decidedly  longer  and  larger  than  the  rest,  the  innermost  odd  one 
being  the  largest  of  the  three,  and  standing  erect  on  the  extreme 
inner  angle  of  the  plate,  and  therefore  nearly  at  right  angles  to  the 
small  spine  within  the  furrow.  The  outermost  spinules  of  these  groups 


VERRILL 

are  similar  in  size  to  those  of  the  adjacent  peractinal  plates,  from 
which  they  are  separated  by  a  distinct  continuous  groove.  The  jaws 
are  covered  with  numerous  erect  spines,  which  are  similar  in  size  and 
form  to  those  of  the  adambulacral  plates,  but  the  adambulacral  plate 
nearest  the  mouth  bears  a  group  of  small  blunt  spinules  deep  within 
the  furrow. 

Eastport,  Maine,  in  shallow  water,  1870  (coll.  A.  E.  Verrill),  type. 
Subsequently  taken  at  several  localities  in  the  Bay  of  Fundy. 

This  variety  is  similar  to  typical  H.  sanguinolenta  in  form  and 
general  appearance,  though  the  dorsal  surface  is  more  uneven  and 
papillose,  owing  to  the  larger  size  of  the  pseudopaxillse  and  the  more 
regular  interspaces;  the  pseudopaxillae  are  generally  more  in  the 
form  of  rounded  fascicles  in  the  latter,  instead  of  regular  comb-like 
groups.  The  differences  are  much  more  marked  on  the  actinal  sur- 
face, where  the  two  regular  rows  of  larger  marginal  plates  and  the 
regular  row  of  peractinal  plates  give  a  very  different  appearance  to 
this  region;  for  in  the  typical  form  these  plates  are  often  scarcely 
distinguishable  in  size,  form,  and  spinulation  from  those  of  the  dorsal 
and  lateral  plates  of  the  rays.  The  adambulacral  and  oral  spines  are 
also  shorter  and  more  crowded  than  in  the  common  form. 

This  form,  which  was  formerly  (1894)  fully  described,  as  a  dis- 
tinct species,  is  of  rather  rare  and  sporadic  occurrence  on  our  coast,  so 
far  as  known.  It  may  be  considered  as  a  vigorous  arctic  or  boreal 
variation  of  the  species,  due,  perhaps,  to  unusually  favorable  condi- 
tions of  climate  and  food.  As  I  have  now  seen  many  specimens  inter- 
mediate between  this  and  the  more  usual  forms  of  the  species,  it  seems 
necessary  to  reduce  it  to  the  rank  of  a  variety  or  subspecies. 

This  subspecies,  like  most  others  of  the  genus,  is  very  variable  in 
the  character  of  the  ossicles  and  spinules,  as  well  as  in  color.  The 
more  common  colors  are  either  orange  or  purple,  rarely  lemon- 
yellow. 

The  type  specimens  of  the  variety  were  from  Eastport,  Maine. 
A  very  small  specimen  has  been  received  from  Popof  Strait 
(W.  H.  Dall,  U.  S.  Nat.  Mus.).  Some  of  those  described  by  Fisher 
appear  to  be  the  same  variety,  especially  his  variety  C.  Some  of  the 
latter  were  very  large ;  radii,  32  mm.  and  235  mm. ;  or  about  nineteen 
inches  in  diameter.  His  variety  C  was  taken  at  eleven  stations  in 
Bering  Sea  and  near  the  Aleutian  Islands,  in  41  to  344  fathoms. 


SHALLOW-WATER    STARFISHES  233 

HENRICIA  SANGUINOLENTA,  Var.  RUDIS  Verrill,  nov. 
Plate  LXXXVI,  figures  5,  50  (details). 

Rays  five,  large,  thick,  tapered  to  small  tips.  Disk  large,  swollen. 
Radii  of  the  type,  22  mm.  and  80  mm. ;  ratio,  i :  3.14;  breadth  of  rays 
at  base,  25  mm.  to  30  mm. 

Dorsal  surface  covered  with  a  multitude  of  very  small,  distinct 
parapaxillse,  forming  a  close  reticulation,  in  which  the  small  papular 
pores  are  mostly  placed  singly.  The  larger  parapaxillae  bear  a  com- 
pact cluster  of  about  five  or  six  slender,  tapered,  acute,  rough  and 
rather  long  spinules ;  many  of  the  smaller  ones  have  three  or  four 
spinules.  These  give  the  surface  a  finely  spinulose  but  rough 
appearance.  There  may  be  six  to  eight  pseudopaxillae  to  a  square 
millimeter. 

The  adambulacral  armature  consists  of  a  single  or  partially  double 
transverse  row,  mostly  of  spatulate  spinules.  There  are  five  to  seven 
larger  spinules  in  each  group,  and  two  or  three  outer  ones  that  are 
much  smaller,  slender,  and  not  flattened.  The  odd  one  on  the  inner 
angle,  which  is  the  largest  one,  is  stout,  wide  at  the  truncate  and 
grooved  tip ;  up  to  2.3  mm.  long ;  the  rest  decrease  gradually  in  size 
and  amount  of  distal  expansion  to  the  most  external  ones.  Adoral 
and  oral  spines  are  similar  in  size  and  form.  The  furrow-spine  is 
single,  small  and  slender. 

The  two  rows  of  marginal  pseudopaxillae  are  small  and  scarcely 
distinguishable  in  the  crowd  of  adjacent  ones,  all  of  which  are  spinu- 
lated  like  the  dorsals.  The  inferomarginals  are  a  little  larger  than 
the  others  and  somewhat  transversely  elongated,  bearing  a  trans- 
verse and  slightly  pectinate  two-rowed  group  of  slender  spinules, 
about  eight  to  twelve  on  the  larger  plates.  The  superodorsals  are 
similar,  when  they  are  distinguishable,  but  in  many  places  they  are 
as  small  as  the  dorso-laterals.  The  peractinals  are  small  and  round- 

»'  TT  *     if  * 

ish.  The  type  is  from  ten  miles  west  of  Point  Franklin,  Arctic 
Ocean,  off  the  north  coast  of  Alaska  in  13^  fathoms,  sand  (coll. 
Murdoch,  Point  Barrow  Expedition,  No.  7623,  U.  S.  Nat.  Mus.). 

This,  though  quite  peculiar  in  respect  to  its  adambulacral  spinules, 
seems  to  be  only  an  extreme  variation  of  H.  sanguinolenta,  perhaps 
only  an  individual  variation.  I  have  seen  no  others  like  it,  though 
some  approach  it  to  some  extent. 


234  VERRILL 

HENRICIA  SANGUINOLENTA  MILIARIS  Verrill,  subsp.  nov. 
Plate  LXXXVIII,  figures  4-40* 

Rays  five,  well  rounded,  evenly  tapered,  and  minutely  spinulated. 
Proportions  about  as  in  the  typical  form  of  H.  sanguinolenta. 
Radii  of  the  type,  10  mm.  and  38  mm. ;  ratio,  1 : 3.8. 

The  surface  appears  to  the  naked  eye  remarkably  fine,  smooth, 
and  even,  owing  to  the  very  small  and  uniform  dorsal  pseudopaxillae. 
These  are  very  numerous,  distinctly  separated,  and  rather  closely 
placed,  but  not  in  contact  nor  forming  rows  or  circles.  They  are 
mostly  roundish,  about  as  high  as  broad,  and  have  about  six  to 
twelve  very  small,  slender,  acute,  rough  spinules.  The  papulae  are 
small  and  numerous  between.  The  dorsal  papulae  are  mostly  single, 
but  often  two  or  three  together  on  an  area. 

On  the  actinal  side,  the  two  marginal  series  and  the  peractinals 
form  three  very  regular  and  evident  rows,  and  there  is  a  very 
evident  channel  between  the  inferomarginals  and  peractinals,  along 
which  there  is  a  very  regular  row  of  single  papulae,  larger  than  those 
found  elsewhere.  Papulae  between  the  peractinals  and  adambulacrals 
are  mostly  lacking.  When  present,  they  are  few  and  very  small. 

The  adambulacral  plates  bear  a  single  rather  slender  furrow-spine, 
and  about  twelve  to  fourteen  on  the  actinal  side,  mostly  in  two 
crowded,  graded  rows.  At  the  edge  of  the  furrow  there  are  three 
larger  than  the  rest,  one  of  which  stands  on  the  apex  of  the  plates ; 
two  others,  nearly  as  large,  stand  just  back  of  the  odd  one,  forming 
an  oblique  pair.  The  rest  are  graded  so  that  the  outer  ones  are 
single  and  of  about  the  same  size  as  the  spinules  of  the  adjacent 
plates.  The  inferomarginals  are  transversely  oblong,  about  twice  as 
broad  as  long,  with  about  fifteen  to  twenty  spinules,  like  those  of  the 
dorsal  ossicles,  but  somewhat  larger.  The  superomarginals  are  about 
half  as  large,  of  similar  form ;  both  rows  are  oblique. 

The  peractinals  are  roundish,  very  distinct,  close  to  the  adambu- 
lacrals, and  have  about  eight  to  ten  spinules  in  a  stellate  group.  On 
some  of  the  rays  they  extend  nearly  or  quite  to  the  tips.  There  is 
usually  a  short,  subactinal  row. 

The  type  was  taken  off  Cape  Cod,  Massachusetts,  in  shallow  water, 
by  the  U.  S.  Fish  Commission,  1879. 

HENRICIA  TUMIDA  Verrill. 
Plate  xn,  figures  i,  2;  plate  LXXXVII,  figures  i,  10  (details). 
Henricia  tumida  VEKRELL,  Amer.  Naturalist,  XLIH,  pp.  554,  555,  fig.  5,  1909. 

Rays  five,  short  and  thick,  acute;  disk  large,  thick,  swollen.  The 
radii  in  the  type  are  1 1  mm.  and  30  mm. ;  in  another,  10  mm.  and 
22  mm. ;  ratios  vary  from  i :  2.00  to  i :  2.7. 


SHALLOW-WATER   STARFISHES  235 

The  dorsal  surface  is  thickly  covered  with  small,  rough  pseudo- 
paxillae,  which  mostly  surround  large  papular  pores,  each  usually 
with  a  single  papula.  The  papular  pores  are  numerous  and  are 
found  on  nearly  all  parts,  even  between  the  peractinals  and  adambu- 
lacral  plates.  Most  of  the  pseudopaxillae  are  roundish  or  elliptical 
and  bear  from  six  to  twelve  spinules;  the  larger  ones  are  narrow- 
oblong,  and  have  about  two  rows  of  spinules,  often  amounting  to 
twenty  to  twenty-five.  Frequently  the  pseudopaxillae  are  crowded 
and  appear  to  blend  together  in  circles  around  the  papulae.  The 
spinules  are  elongated,  slender,  with  rough,  thorny  or  spiculose  tips. 

The  adambulacral  plates  and  spines  are  relatively  large;  each 
plate  bears  a  rather  short  intra-ambulacral  or  furrow-spine,  and  a 
transverse  row  of  four  to  six  on  the  actinal  side.  The  first  three  of 
these  are  larger  and  divergent,  elongated,  and  obtuse.  The  others 
are  smaller  and  graded. 

The  inferomarginals  form  regular  rows;  they  are  strongly  com- 
pressed and  bear  about  twelve  to  twenty  small,  slender  spinules. 
The  superomarginals  are  smaller,  oblong,  oblique,  compressed,  with 
similar  but  fewer  spinules,  like  those  of  the  dorsals.  The  peractinals 
are  small  and  extend  to  about  the  distal  fourth.  They  have  four  to 
six  spinules ;  they  are  close  to  the  adambulacrals ;  one  or  two  very 
short  intermarginal  rows,  or  a  small  group,  may  be  present  close  to 
the  base  of  the  ray.  They  are  similar  to  the  superomarginals  and 
dorsals  in  spinulation. 

The  central  part  of  the  disk  and  oral  region  are  deeply  withdrawn, 
forming  a  funnel-like  pit.  The  interradial  actinal  areas  are  closely 
folded  inward,  and  about  eight  or  nine  adambulacral  plates  in  each 
row  are  included  in  the  incurved  portion  of  the  actinal  surface. 
These  plates  and  their  spines  rapidly  decrease  in  size  adorally,  those 
near  the  jaws  being  less  than  half  as  large  as  the  eighth  or  ninth. 
There  are  two  short,  stout,  apical  peroral  jaw-spines,  and  about  six 
small  epiorals. 

This  species  or  "  form "  is  easily  distinguished  by  its  large, 
swollen  disk  and  short  rays,  and  by  the  deeply  sunken  oral  region. 

The  dorsal  surface  is  nearly  as  in  H.  borealis,  except  that  the 
ossicles  are  usually  smaller  and  the  spinules  are  more  numerous  and 
in  smaller  clusters,  and  often  stand  in  small,  rounded  or  oblong 
groups  around  the  papulae. 

The  marginal,  actinal,  and  adambulacral  plates  and  spines  are 
nearly  the  same  as  described  under  H.  borealis,  but  in  some  speci- 
mens the  supermarginal  row  of  plates  is  not  separated,  proxi- 


236  VERRILL 

mally,  from  the  lower  row  by  interpolated  plates  and  is,  therefore, 
more  regular  and  distinct.  Its  broad  disk  and  very  short  rays  are 
shared  by  H.  arctica;  but  in  the  latter  the  disk  is  flat  and  the  oral 
region  is  not  sunken,  while  the  spinulation  is  very  minute,  and  the 
spinules  far  more  numerous  and  differently  arranged  on  all  the 
plates,  but  particularly  on  the  adambulacrals. 

Dutch  Harbor,  Unalaska  (Harriman  Expedition,  Dr.  W.  R. 
Coe). 

The  name  tumida  was  originally  a  manuscript  name  used  on  labels 
by  Dr.  Richard  Rathbun,  who  had  recognized  this  as  a  distinct  form 
several  years  ago. 

This  is  a  very  peculiar  form  of  Henrida,  having  a  large,  swollen 
disk  and  very  short  rays,  but  in  most  other  respects  agreeing  so 
closely  with  H.  borealis  that  it  seems  possible  that  it  is  only  a  sexual 
form  or  a  variation  of  the  latter. 

The  peculiar  excavate  condition  of  the  actinal  surface  of  the  disk 
and  oral  area  seems  to  be  specially  adapted  to  the  carrying  of  the 
eggs  and  young.  Whether  all  the  specimens  were  actually  females 
could  not  be  determined  from  dry  specimens.  It  might  be  thought 
that  this  is  the  normal  female,  while  H.  borealis  is  the  male  of  the 
same  species,  were  it  not  that  Dr.  Fisher  has  figured  the  female  of 
H.  borealis,  while  actually  carrying  eggs,  and  it  does  not  agree  with 
H.  tumida,  but  has  about  the  proportions  of  borealis,  and  arches  its 
disk  as  is  usual  in  other  species  of  the  genus.  There  is  no  evidence 
given  that  the  arching  is  persistent,  nor  that  the  adoral  plates  and 
spines  are  modified  for  that  purpose. 

HENRICIA  TUMIDA  BOREALIS  Verrill,  subsp.,  nov. 

Plate  xn,  figures  3,  4;  plate  LXXXVI,  figures  6-60  (details)  ;  plate  LXXXVIII, 
figures  5-5b   (details). 

Henrida  eschrichtii  (pars}  FISHER,  op.  cit.,  191  ib,  p.  276,  pi.  LXVII;  pi.  Lxvin, 
fig.  I  (non  Mtiller  and  Troschel). 

Rays  five,  short  or  moderately  long,  stout,  swollen  at  base,  taper- 
ing rapidly  to  slender  tips.  Disk  rather  large  and  thick.  Radii  of  a 
dry  specimen,  12  mm.  and  42  mm. ;  ratio,  about  i :  3.5.  Breadth  of 
rays  at  base,  15  mm.  Another  specimen  has  the  radii  u  mm.  and 
42  mm. ;  ratio,  about  i :  3.8.  A  short-rayed  specimen  has  the  radii 
10  mm.  and  24  mm. ;  ratio,  i :  2.4. 

The  dorsal  surface,  in  dry  specimens,  has  a  distinctly  areolate 
appearance,  the  angular  meshes  of  the  network  of  ossicles  being 
larger  and  deeper  than  usual  in  the  related  species,  and  the  narrow 


SHALLOW-WATER   STARFISHES  237 

ossicles  everywhere  covered  with  even,  fine,  slender,  upright, 
crowded  spinules,  which  largely  stand  in  two  or  three  close  rows  on 
the  larger  ossicles,  and  sometimes  in  single,  comb-like  rows,  around 
and  between  the  papulae.  The  larger  groups  are  often  curved  or 
crescent-shaped.  These  spinules  are  longer  and  more  slender  than 
in  ordinary  H.  sanguinolenta,  and  have  rough,  acute,  or  subacute 
tips.  They  diverge  but  little.  The  papular  pores  are  rather  large 
and  mostly  isolated.  The  madreporic  plate  is  of  fair  size,  rough,  with 
rows  of  small  spinules  on  the  gyri. 

The  dorsal  ossicles,  when  the  spines  are  removed,  are  small, 
thick,  convex,  often  narrow-oblong.  They  form  an  open  reticulation 
in  which  the  smaller  meshes  are  polygonal  or  irregularly  triangular. 
The  upper  and  lower  marginal  plates  are  easily  distinguished  by 
their  larger  size  and  more  convex  surface.  The  two  rows  are  con- 
tingent on  the  distal  third  of  the  rays,  with  large  papular  pores 
between  them.  They  become  separated  by  a  row  of  small  rounded 
ossicles,  increasing  in  size  proximally.  Two  or  three  other  rows  of 
small,  irregular  intermarginal  ossicles  are  also  interpolated  at  the 
base  of  the  rays,  in  the  lateral  interradial  areas,  where  the  upper  and 
lower  marginals  are  widely  separated.  The  superomarginals  are 
prominent,  with  a  narrow  transversely  oblong  surface.  The  infero- 
marginals  are  larger,  with  the  exposed  surface  sometimes  angular, 
or  transversely  oblong,  sometimes  curved  or  crescent-shaped,  with  a 
narrow,  prominent  ridge  for  the  insertion  of  the  spines.  They  are 
always  decidedly  longer  transversely  to  the  rays ;  the  transverse 
diameter  is  often  double  the  longitudinal.  When  curved,  the  con- 
vexity is  toward  the  base  of  the  ray.  Some  of  the  proximal  ones 
are  much  compressed,  with  a  rather  sharp  ridge,  and  bear  only  one 
row  or  comb  of  spinules. 

The  peractinal  ossicles  form  but  one  row,  though  often  the  row 
is  irregular.  They  are  somewhat  rounded  or  lobate  lozenge-shaped, 
and  much  smaller  transversely  than  the  inferomarginals,  but  equal 
to  them  in  number.  The  two  rows  are  united  by  apophyses,  between 
which  is  a  row  of  large  papular  pores.  They  are  united  directly  to 
the  adambulacral  ossicles,  with  which  they  agree  in  length.  The 
latter  are  transversely  oblong  and  prominent. 

The  spinules  on  the  superomarginal  and  lateral  ossicles  are  about 
the  same  as  those  of  the  dorsal  surface,  but  on  the  inferomarginal 
plates  they  become  a  little  larger  and  longer.  On  the  inferomarginals 
they  mostly  stand  in  two  rows,  of  about  eight  to  sixteen  each,  along 
the  middle  of  the  rays ;  but  beneath  the  disk  they  may  be  reduced  to 


238  VERRILL 

a  single  comb-like  row  of  six  to  twelve,  on  some  of  the  plates.  The 
peractinal  plates  bear  a  smaller  double  group  of  similar  spines,  mostly 
six  to  ten  in  number.  The  spines  on  the  actinal  surface  of  the  adam- 
bulacral  plates  stand  in  two  rows,  or  often  only  one  row  distally, 
increasing  regularly  in  size  and  length  to  the  one  or  two  unpaired 
ones  at  the  edge  of  the  groove.  These  are  rather  large,  slightly 
clavate,  and  obtuse.  The  intra-ambulacral  spine  is  much  smaller  and 
shorter,  acute  or  subacute.  not  very  slender.  The  two  apical  peroral 
spines  are  rather  stout,  subacute ;  the  epiorals  and  adorals  are  similar. 

Taken  at  Dutch  Harbor,  Unalaska,  Yakutat,  Fox  Cape,  Sitka,  etc. 
(Harriman  Expedition).  Common. 

Recorded  by  Dr.  Fisher  (as  eschrichtii)  from  numerous  localities 
between  Bering  Straits  and  Yakutat,  Alaska,  from  the  shore  to  86 
fathoms,  and  on  the  Asiatic  side  to  Kamchatka  and  the  Kuril  Islands. 

Dr.  Fisher  records  specimens  of  this  species  carrying  eggs,  taken 
June  8,  10,  and  14,  1906;  and  one  carrying  young  up  to  4  mm.  in 
diameter  with  eight  pairs  of  ambulacral  feet  to  each  ray;  taken  by 
the  "  Corwin,"  in  Bering  Sea.  (See  Fisher's  pi.  LXVIII,  fig.  i.) 

Probably  this  subspecies  has  formerly  been  confused  with  H.  san- 
guinolenta, which  it  much  resembles,  superficially.  > 

It  can  best  be  distinguished  from  H.  sanguinolenta  by  the  larger 
and  decidedly  transversely  oblong  inferomarginal  ossicles  and  their 
clusters  of  spines,  of  the  same  form.  These  plates  in  H.  sanguino- 
lenta are  usually  smaller,  scarcely  oblong,  and  bear  smaller  divergent 
clusters  of  spines.  This  is  also,  in  most  cases,  a  stouter  species,  with 
shorter  rays  and  much  larger  disk.  Other  differences  appear  on 
closer  examination,  especially  in  the  dorsal  skeleton  and  marginal 
plates,  and  in  the  much  narrower  and  simpler  rows  and  longer  adam- 
bulacral  spines. 

It  is  so  unlike  H.  leviuscula  that  there  is  no  need  to  compare  the 
two,  in  detail. 

This  subspecies  is,  apparently,  quite  unlike  H.  eschrichtii  M  tiller 
and  Troschel,  as  already  remarked.  The  latter  was  described  as 
having  the  proportions  1 : 4,  or  about  those  of  ordinary  specimens  of 
typical  H.  sanguinolenta,  from  which  it  differs  less  than  var.  pec- 
tinata,  or  subsp.  miliaris.  I  consider  it  nearly  the  same  as  var.  pec- 
tinata. 

One  specimen,  taken  with  the  type  of  H.  tumida,  has  nearly  the 
same  spinulations  as  the  latter,  both  above  and  below,  and  yet  does 
not  have  the  oral  region  inarched.  It  has,  however,  a  somewhat 
smaller  and  less  turgid  disk.  Its  radii  are  8  mm.  and  20  mm. ;  ratio, 
1:2.5. 


SHALLOW-WATER   STARFISHES  239 

Some  of  the  varietal  forms  referred  to  H.  sanguinolenta  by  Fisher 
appear  to  me  to  belong  rather  with  borealis,  especially  those  on 

pi.  LXVIII,  fig.  3. 

It  is  not  unlikely  that  H.  borealis  may  be  the  male  form  of  H. 
tumida.  Should  that  prove  to  be  the  case,  the  latter  name  has 
priority. 

HENRICIA  ARCTICA  Verrill,  sp.  nov. 
Plate  LXXXVII,  figures  3-3c  (details)  ;  text-figure  No.  13. 

Henricia  eschrichtii  (pars')   FISHER,  op.  cit,  19116,  p.  276,  pi.  LXVIII,  fig.  2 
(non  Muller  and  Troschel). 

Form  stellate,  with  a  wide  disk  and  short,  thick  rays.  Radii  of 
the  type,  13  mm.  and  28  mm.;  ratio,  1:2.15;  breadth  of  rays  at 
base,  15  mm. 


FIG.  13. 

Henricia  arctica  Ver.,  type.     U.  S.  Nat.  Mus.     Actinal  side,  natural  size. 

The  disk,  as  preserved,  is  but  little  convex  dorsally,  and  nearly 
flat  on  the  actinal  side,  with  no  marked  oral  concavity,  the  jaws  and 
oral  spines  being  fully  exposed,  not  sunken  in  a  deep  concavity  as 
in  H .  tumida. 

The  dorsal  ossicles  are  very  small  and  closely  united  into  a  very 
fine-meshed  reticulation,  with  small,  single  papular  pores.  The  small 
pseudopaxillae  are  covered  with  compact  groups  of  very  small,  short 
spinules  of  nearly  uniform  length,  ten  to  fifteen  or  more  on  the 
larger  ones,  often  only  four  to  six  on  the  smaller  ones.  They  pro- 
duce a  pretty  even,  almost  granular-appearing  surface,  of  fine 
texture.  Very  often  four  to  six  small  pseudopaxillae  form  a  rosette- 
like  group  around  a  papular  pore. 


240  VERRILL 

The  spinules  are  smaller  and  shorter  than  in  any  other  North 
Pacific  or  Arctic  species  known  to  me,  and  the  surface  is  corre- 
spondingly more  even  and  finer-grained. 

The  two  marginal  rows  and  the  peractinal  row  of  plates  are  dis- 
tinct, but  not  conspicuous  and  not  very  regular.  The  inferomar- 
ginals  are  larger  than  those  of  the  other  rows.  They  are  obliquely 
placed  and  transversely  compressed  proximally,  but  become  more 
elliptical  or  rounded  and  more  irregular  distally.  They  bear  a  close 
cluster  of  minute  spinules  like  those  of  the  dorsals.  The  infero- 
marginals  are  smaller  and  scarcely  distinguishable  in  many  places. 
There  are  two  or  three  short  rows  of  mtermarginals  proximally. 
The  peractinal  plates  are  pretty  regular,  oblique,  mostly  rhombic, 
smaller  than  the  inferomarginals,  and  spinulated  in  the  same  way. 

The  adambulacrals  bear  several  close  irregular  rows  of  very  short, 
and  very  small,  obtuse  spinules,  about  five  to  eight  in  each  row,1  not 
very  unlike  in  size,  and  an  odd  one  slightly  larger  on  the  inner  angle ; 
the  furrow-spine  is  very  small  and  short.  The  adoral  and  oral  spines 
are  larger,  obtuse,  numerous  and  crowded. 

This  species  appears  to  be  more  distinct  from  all  the  other  North 
Pacific  forms  of  the  genus  than  most  of  those  hitherto  described,  at 
least  from  the  shallower  waters. 

Why  Dr.  Fisher  should  have  united  it  with  his  H.  eschrichtii  is 
not  easy  to  understand,  for  there  is  no  particular  resemblance  to  that 
species,  unless  in  the  short  rays;  but  in  this  respect  the  present 
species  exceeds  all  others  except  H.  tumida. 

From  the  latter  it  differs  widely  in  the  much  finer  and  shorter 
dorsal  and  marginal  spinules,  in  the  smaller  and  closer  ossicles,  and 
especially  in  the  very  different  character  of  the  adambulacral 
spinules,  which  are  much  longer  and  fewer  in  H.  tumida,  and  in  a 
single  row.  The  latter,  moreover,  has  a  deep  concavity  around  the 
oral  region,  in  which  the  jaws  and  oral  plates  are  so  deeply  sunken 
as  to  be  seen  with  difficulty.  Possibly  this  last  character  pertains 
only  to  the  female,  as  suggested  above,  but  in  that  case  the  other 
characters  named  are  ample  for  distinguishing  the  two  forms. 

The  type  of  this  species  was  from  off  Cape  Lisburne,  Alaska  (coll. 
H.  D.  Woolfe,  No.  12,818,  U.  S.  Nat.  Mus.).  The  specimen  figured 
by  Fisher  is  from  the  same  vicinity. 

1  The  figure  (pi.  LXXXVII,  fig.  30)  represents  these  spines  as  too  few  and  too 
large.  In  the  specimen  figured  the  spines  were  badly  worn  off  from  most  of 
these  plates. 


SHALLOW-WATER   STARFISHES  24! 

HENRICIA  LONGISPINA  Fisher. 

Henricia  longispina  FISHER,  op.  cit.,  1910,  p.  572;  19116,  p.  299,  pi.  LXXVI, 
figs.  I,  2 ;  pi.  cxi,  figs.  3,  30. 

Rays  five,  moderately  slender,  cylindrical.  Radii  of  type,  9  mm. 
and  47  mm. ;  ratio,  1 :  5.2.  Dorsal  ossicles  are  stout  and  form  a 
rather  coarse  and  open  reticulation.  Spinules  are  spaced  in  fascicu- 
late groups  of  two  to  nine,  mostly  five  to  seven,  unusually  long  for 
the  genus,  up  to  I  mm.  to  1.5  mm.,  acute  and  finely  denticulate. 
Marginal  rows  of  plates  fairly  regular,  not  very  large,  each  with  six 
to  nine  spinules.  The  inferomarginal  spines  are  a  little  the  larger; 
somewhat  transversely  elongated;  two  or  three  intermarginal  rows 
of  small  plates  proximally,  the  longest  extending  to  one-half  to  two- 
thirds  the  length  of  the  ray;  interradial  plates  small,  extending  to 
about  half  the  length  of  the  ray,  with  two  to  four  spinules.  Papulae 
exist  between  the  interactinal  and  adambulacral  plates,  as  in  most 
species  of  the  genus.  Adambulacral  plates  have  one  saber-shaped 
furrow-spine;  on  the  actinal  margin  a  long,  tapered  spinule  with 
two  smaller  ones  seated  back  of  it,  followed  by  three  or  four  smaller 
and  shorter,  acute,  graded  spinules,  the  whole  series  forming  a 
zigzag  or  irregular  transverse  row.  The  adambulacral  spines  are 
longer  than  any  others. 

Color  in  life,  milk  white. 

The  type  was  from  Queen  Charlotte  Sound,  off  Vancouver  Island, 
in  68  to  107  fathoms,  soft  mud.  The  only  other  locality  is  Naha 
Bay,  Behm  Canal,  southern  Alaska,  in  41  to  134  fathoms,  gravel. 

The  description  is  abridged  from  that  of  Professor  Fisher. 

HENRICIA  ASPERA  Fisher. 
Henricia  aspera  FISHER,  op.  cit.,  19116,  p.  293,^!.  LXXV,  figs.  1-5. 

Stellate,  usually  with  five  long,  slender  rays.  Radii  of  type, 
15  mm.  and  100  mm.;  ratio,  about  1:6.6;  sometimes  1:7.2,  and 

i :  5-3- 

Dorsal  plates  openly  reticulated,  the  spinules  not  grouped  into 
evident  pseudopaxillae,  but  short,  granule-like  and  nearly  continuous 
over  the  lines  of  ossicles  around  the  papular  pores;  the  meshes  and 
papular  areas  rather  large,  each  with  several,  five  to  twelve,  papulae 
in  adults.  Spinules  stout,  sharp,  not  crowded,  often  obscured  by 
membrane;  the  rows  often  interrupted. 

Marginal  plates  in  regular  rows,  the  upper  ones  usually  a  little 
smaller,  spinules  about  ten  to  twelve  to  a  plate.  Interactinal  plates 

17 


242  VERRILL 

in  two  rows  proximally.  Adambulacral  plates  have  one  or  two 
furrow-spines  proximally ;  on  actinal  side  two  larger  stoutish  spines 
obliquely  placed;  and  outside  these,  three  or  four  smaller  graded 
spines  in  an  irregular  transverse  row. 

Mr.  Fisher  records  this  from  numerous  localities  in  26  to  313 
fathoms,  from  Bering  Sea  to  Santa  Barbara,  California. 

The  above  description  is  condensed  from  Fisher. 

Family  SOLASTERID^E  Perrier. 

Solasterina  (sub-family  of  Echinasteridae)  VIGUIER,  Squelette  des  Stellerides, 

Nouv.  Arch.  Zool.  Exper.,  vn,  p.  133,  pi.  vnr,  figs.  1-7,  1878  (structure). 
Solasteridce  PERKIER,  Etoiles  de  mer,  p.  210,  1884.     Sladen,  Rep.  Voy.  Chall., 

Zool.,  xxx,  p.  442,  1889.     Perrier,  Exped.  Trav.  et  Talism.,  p.  151,  1894. 

Fisher,  op.  cit,  191  ib,  p.  305. 
Echinasteridce  (Pars)  PERRIER,  Revis.  Stell.,  Nouv.  Arch,  du  Mus.,  iv,  p.  358, 

1875- 

Rays  varying  in  number  from  five  to  thirteen  or  more.  Dorsal 
skeleton  generally  formed  either  by  openly  reticulated  or  loosely 
imbricated  (rarely  detached,  L&tmaster)  ossicles,  which  have  a  con- 
vex or  elevated  central  boss,  bearing  a  cluster  of  slender,  paxilliform, 
movable  spinules,  webbed  together  into  a  stellate  or  penicillate  group 
(pseudopaxillae).  Both  series  of  marginal  plates  are  usually  dis- 
tinguishable, at  least  distally,  alternate  or  opposite,  or  in  one  line. 
The  upper  ones  are  often  the  smaller  and  essentially  like  the  dorsal 
pseudopaxillae.  Both  rows  bear  paxilliform  spinelets. 

Actinal  interradial  pseudopaxillae  are  generally  present  and  similar 
to  the  dorsals.  Adambulacral  plates  are  transversely  elongated  and 
bear,  each,  one  to  five  or  more  spines,  often  webbed,  in  a  longitudinal 
row  on  the  inner  margin  of  the  groove,  and  one  or  more  clusters  or  a 
transverse  webbed  series  or  comb  of  movable  spines  on  the  outer 
surface.  True  pedicellariae  are  not  known  in  most  species;  but 
large  specimens  sometimes  have  a  few  small,  bifid  spines  in  the 
furrow  series,  and  in  S.  stimpsoni  a  few  minute  bivalve  pedicellariae 
sometimes  occur  on  the  dorsal  paxillar  areas.  Tube-feet  have 
suckers. 

Sladen  (1888  and  1889)  divided  this  family  into  two  subfamilies: 
Solasterinae  and  Korethrasterinae.  In  this  he  was  followed  by  Perrier 
(1894,  pp.  154,  158).  Later  these  have  been  considered  separate 
families. 

Genus  Solaster  Forbes. 

Solaster  (pars)   FORBES,  Mem.  Wern.  Soc.,  vni,  p.  120,  1839;  British  Star- 
fishes, p.  109,  1841.    Gray,  Ann.  and  Mag.  Nat  Hist.,  vi,  p.  183,  November, 


SHALLOW-WATER   STARFISHES  243 

1840;  Synopsis,  p.  4,  1866.  VERRILL,  Proc.  Boston  Soc.  Nat.  Hist,  x, 
P-  345.  !866  (Solaster  and  Crossaster  first  separated).  Agassiz,  North 
American  Starfishes,  pp.  m,  112,  1887  (structure).  Perrier,  Exped. 
Trav.  et  Talism.,  p.  154,  1894.  Sladen,  op.  cit,  p.  450,  189. 

Crossaster  (pars)  MULLER  and  TROSCHEL,  Monatsb.  Preuss.  Akad.  Wiss., 
Berlin,  April,  1840,  p.  103. 

Solaster  (pars')  VIGUIER,  Nouv.  Arch.  Zool.  Exper.,  vii,  p.  138,  1878  (struc- 
ture). Bell,  Catal.  Brit.  Echinod.,  p.  88,  1892. 

Solaster  (pars)  DANIELSSEN  and  KOREN,  Norw.  N.  Atlantic  Exp.,  Asteroidea, 
pp.  42,  52,  53,  1884  (structure).  Fisher,  op.  cit.,  ignb,  306. 

Disk  rather  broad.  Rays  seven  to  twelve,  variable  in  each  species. 
Dorsal  ossicles  small,  mostly  subcruciform  or  slightly  four-lobed, 
sometimes  stellate,  usually  reticulated,  or  in  quincunx,  on  the  disk 
and  proximal  part  of  the  rays,  but  closely  imbricated  near  ends  of  the 
rays,  and  often  arranged  in  regular  quincunx  order;  sometimes 
united  by  smaller  ossicles.  They  are  convex  or  slightly  mammillate, 
with  a  central  boss,  and  bear  a  stellate  or  fasciculate  cluster  of 
slender  spinules,  webbed  together,  and  usually  enclosing  one  or 
several  central  spinules,  connected  by  the  web,  forming  pseudo- 
paxillae.  The  outer  circle  of  webbed  spinules  often  forms  a  funnel- 
shaped  structure  in  life,  or  when  well  preserved;  but  the  spinules 
are  movable,  and  in  dry  specimens  they  are  usually  mostly  collapsed 
or  tipped  over,  thus  forming  irregular  groups  or  pencils  or  small 
spinules,  from  five  to  twenty  or  more  in  a  group. 

The  superomarginal  plates  are  small  and  close  to  the  larger  infero- 
marginals,  usually  alternating.  They  are  usually  hardly  distinguish- 
able from  the  dorsal  pseudopaxillae,  except  near  the  ends  of  the  rays, 
but  they  extend  regularly  to  the  apical  plate. 

Inferomarginals  much  larger  and  more  elevated,  transversely 
oblong,  forming  a  rather  conspicuous  row.  They  bear  a  large  num- 
ber of  paxilliform  spinules  in  two  or  more  transverse  rows.  Adam- 
bulacral  plates  usually  have  two  to  four  shorter  groove-spines, 
webbed  together,  and  an  actinal  transverse  row  of  four  to  nine  or 
more  longer  spines,  also  webbed. 

The  interradial  actinal  areas  are  small,  but  distinct,  and  bear 
pseudopaxillae,  much  like  the  dorsal  ones. 

Papulae  are  numerous  on  the  dorsal  surface,  but  stand  singly  or  in 
small  groups.  A  single  row  of  peractinal  pseudopaxillae  usually 
extends  along  the  proximal  part  of  the  rays,  to  about  the  middle. 

The  eggs  and  young  are  carried  attached  to  the  oral  region  in 
clusters  in  5".  endeca  and  other  species. 

The  structure  of  the  skeleton  has  been  very  fully  described  and 
beautifully  illustrated  by  Agassiz  in  the  work  cited  above. 


244  VERRILL 

The  species  of  this  genus,  like  those  of  Henricia  and  Leptasterias, 
are  numerous  and  variable,  presenting  in  many  cases  puzzling  forms, 
due,  perhaps,  to  hybridism  or  to  the  localization  of  accidental 
varieties  or  "  sports,"  and  made  possible  by  the  habit  of  the  parents 
to  carry  and  care  for  the  eggs  and  young,  which  have  no  free- 
swimming  stages.  (See  remarks  under  Leptasterias,  pp.  116,  117.) 

SOLASTER  ENDECA  (Linne)  Forbes. 

Plate  ix,  figures  2,  3   (young)  ;  plate  LXXXVII,  figures  4-46   (details)  :  plate 
LXXXIX,  figure  i  (typical). 

Asterias  endeca  LJNN£,  Mant.  Plant.  App.,  p.  543,  1771.  Retzius,  K.  Svenska 
Vet.  Akad.  Handl.,  iv,  p.  237,  1783.  Gmelin,  Syst  Nat.,  p.  3162,  1788. 
Lamarck,  Anim.  s.  Verteb.,  n,  p.  560,  1816. 

Solaster  endeca  FORBES,  Mem.  Wern.  Soc.,  vni,  p.  121,  1839;  Hist.  British 
Starfishes,  p.  109  (fig.),  1841.  Mtiller  and  Troschel,  Syst,  p.  26,  1842. 
Gray,  Ann.  and  Mag.,  p.  183,  1840;  p.  19,  1848;  Synopsis,  p.  5,  1866. 
Stimpson,  Invert  Grand  Manan,  p.  14,  1853.  Lutken,  Oversigt  over 
Gronlands  Echinodermata,  p.  35,  1857.  Verrill,  Proc.  Boston  Soc.  Nat. 
Hist,  x,  pp.  345,  356,  1866;  Bull.  Essex  Inst,  in,  p.  4,  1871;  Amer. 
Journ.  Sci.,  v,  p.  104,  1873;  Expl.  of  Casco  Bay,  p.  356,  1874.  Perrier, 
Stellerides  du  Mus.,  in  Arch.  Zool.  Exper.,  TV,  p.  359,  1875.  A.  Agassiz, 
N.  Amer.  Starfishes,  p.  112,  pi.  xvn,  figs.  1-5,  1877.  Viguier,  Squelette 
des  Stellerides,  Arch.  d.  Zool.  Exp.  et  Gen.,  vn,  p.  134,  1878.  Duncan 
and  Sladen,  Echinod.  Arctic  Sea,  p.  40,  pi.  in,  figs.  5-8,  1881.  Danielssen 
and  Koren,  Norske  Nordshav.  Exp.,  xi,  p.  50,  pi.  ix,  fig.  13,  1884.  Verrill, 
Results  Expl.  by  Albatross  in  1883,  p.  541,  1885.  Murdoch,  op.  cit,  1885,  p. 
160.  Ganong,  Echinod.  New  Brunswick,  p.  33,  pi.  i,  fig.  8,  1888.  Fewkes, 
Bull.  Essex  Inst,  xxin,  p.  63,  1891.  Bell,  Catal.  British  Echinod.,  p.  90, 
1892.  Verrill,  Amer.  Journ.  Sci.,  xux,  p.  199,  1895  (distribution).  Lud- 
wig,  Fauna  Arctica,  I,  p.  464,  1900  (synonymy  and  distribution).  Clark, 
op.  cit,  1904,  p.  556,  pi.  HI,  figs.  12,  13;  pi.  iv,  fig.  23.  Fisher,  op  cit.,  191  ib, 
P-  3O7,  pi.  LXXXI  ;  pi.  LXXXII,  figs,  i,  2,  4. 

A  very  large  ten-rayed,  typical  specimen,  collected  by  Mr.  N.  P. 
Scudder,  off  the  coast  of  Greenland  (lot  61),  has  the  following 
characters : 

Radii,  76  mm.  and  190  mm. ;  ratio,  nearly  as  i :  2.5.  The  adambu- 
lacral  plates,  opposite  the  bases  of  the  rays,  mostly  bear  two  unequal, 
short  furrow-spines,  and  a  smaller  rudimentary  or  pedicellaria-like 
spine ;  on  the  ventral  side  they  bear  an  obliquely  transverse  group  of 
eight  to  fifteen  longer  and  stouter  graded  spines,  of  which  the  inner 
ones  are  longer,  the  first  two  being  considerably  longer  and  placed 
nearly  side  by  side.  They  are  partially  webbed  together  and  often 
stand  more  or  less  in  two  alternate  rows.  Outside  each  of  these, 
but  often  almost  blending  with  them,  there  is  a  group  of  fifteen  to 


SHALLOW-WATER   STARFISHES  245 

twenty  or  more  smaller  actinal  spinules,  standing  irregularly  in 
about  three  transverse  rows  on  each  peractinal  plate.  Farther  out 
they  rapidly  decrease  in  number  and  disappear  at  about  the  middle 
of  the  free  part  of  the  ray.  Outside  these  there  is  a  conspicuous  row 
of  inferomarginal  pseudopaxillae,  distinctly  larger  than  the  others. 
Each  of  these  plates  bears  about  thirty  to  forty  small,  slender,  rough, 
subacute  spinules,  in  three  or  four  divergent  rows.  The  actinal 
inter  radial  plates  bear  clusters  of  spinules  (pseudopaxillae)  similar  to 
those  of  the  lower  marginals,  but  a  little  smaller  and  more  com- 
pressed, and  mostly  webbed  together  laterally.  They  are  radially 
arranged  and  rather  numerous,  forming  about  sixteen  to  twenty 
rows,  covering  a  rather  large  acute-triangular  interradial  area. 

The  superomarginal  groups  are  much  smaller,  but  prominent  and 
unequal  in  size;  the  largest  are  ovate  or  oblong-ovate,  bearing 
twenty  to  thirty  small  acute  spinules  in  about  three  or  four  divergent 
rows.  They  are  close  to  the  inferomarginal  ones  and  alternate  with 
them.  The  dorsal  pseudopaxillae  are  rather  small  and  unequal  in 
size ;  the  larger  are  oblong-elliptical ;  the  smaller  oval  or  round ;  all 
are  covered  with  minute  divergent  spinules  of  nearly  uniform 
length.  The  smaller  plates  bear  about  seven  to  nine  spinules;  the 
larger  fifteen  to  twenty  or  more.  When  expanded  they  are  stellate 
and  webbed  together,  with  two  or  three  in  the  middle.  Madreporic 
plate  large  and  irregular,  with  somewhat  enlarged  and  prominent 
pseudopaxillae  surrounding  it.  The  papulae  are  large  and  numerous ; 
they  mostly  stand  singly  or  two  together  in  the  interspaces  between 
the  plates. 

Greatest  diameter,  15  inches,  or  382  mm.  Radii  of  disk,  3  to  3.5 
inches,  or  76  mm.  to  89  mm.  Radii  of  rays,  7  to  7.5  inches,  or 
178  mm.  to  190  mm.  Breadth  of  rays,  at  base  2  inches,  or  51  mm. 

Another  very  similar  specimen  from  the  Fishing  Banks  off  Nova 
Scotia  (lot  820)  has  nine  rays.  Its  dimensions  are  as  follows : 

Greatest  diameter,  13.50  inches,  or  344  mm.  Radii  of  disk,  2.75 
to  3  inches,  or  70  mm.  to  76  mm.  Radii  of  rays,  6.50  to  6.75  inches, 
or  166  mm.  to  174  mm.  Breadth  of  rays  at  base,  1.75  inches,  or 
44  mm.  Ratios  of  radii,  about  1 : 2.5. 

The  dorsal  pseudopaxillae  are  so  close  that  when  the  spinules  are 
fully  spread  out  or  naturally  expanded  in  the  stellate  and  webbed 
condition,  they  nearly  or  quite  touch  each  other,  leaving  room  only 
for  the  mostly  solitary  papulae  between  them.  They  are  nearly  uni- 
form in  character  over  the  disk  and  rays,  though  variable  in  size. 
On  the  sides  of  the  rays  proximally  they  are  arranged  pretty 


246  VERRILL 

regularly  in  alternating  obliquely  transverse  rows,  or  in  quincunx, 
and  are  a  little  larger  and  bear  twelve  to  eighteen  or  more  divergent 
webbed  spinules  around  the  edge  and  four  to  six  or  more  on  the 
central  part.  On  the  distal  part  of  the  rays  the  ossicles  become 
smaller,  more  closely  crowded,  and  imbricated,  with  their  regularly 
stellate  spinules  in  contact.  Many  of  these  have  only  nine  to  twelve 
spinules  around  the  edge  and  one  to  three  in  the  middle. 

The  adambulacral  plates  mostly  bear  two  unequal  furrow-spines, 
but  some  proximal  ones  bear  three,  the  central  much  the  larger ;  dis- 
tally  they  mostly  have  but  one. 

NORTH  PACIFIC  SPECIMENS. 

A  ten-rayed  specimen  taken  off  Kadiak,  Alaska,  in  5  fathoms,  by 
the  Harriman  Expedition,  agrees  well  with  New  England  specimens 
of  the  same  size.  Its  color  in  life  was  "  orange  with  purple  bars  " 
(Coe). 

Radii,  25  mm.  and  65  mm. ;  ratios,  about  1 : 2.6.  The  rays  are 
convex  above  and  taper  to  rather  slender  tips.  Dorsal  surface 
closely  and  evenly  covered  with  small,  stellate  pseudopaxillae,  which 
do  not  form  any  distinct  radial  rows,  but  often  are  in  contact  by 
their  edges.  When  expanded  they  are  unequal,  mostly  circular  and 
regularly  stellate,  each  having  from  nine  to  fifteen  or  more  minute, 
divergent  spinules,  webbed  together  around  the  edge,  and  two  to 
four  in  the  middle.  They  are  borne  on  very  small,  convex,  mostly 
four-lobed  ossicles,  between  which  are  numerous  isolated  papular 
pores,  each  of  the  larger  pseudopaxillae  having  three  to  five  around 
it.  On  the  sides  of  the  rays  the  pseudopaxillae  are  in  pretty  regular 
quincunx  order. 

Central  (nephridial)  pore  distinct.  M^dreporic  plate  large,  with 
several  pseudopaxillae  on  its  surface. 

Superomarginal  pseudopaxillae  small,  but  more  than  twice  as 
large  as  the  dorsals,  with  numerous  larger,  closely  clustered  spinules. 
They  are  round  or  ovate,  close  to  the  upper  side  of  the  inferomar- 
ginals,  with  which  they  alternate. 

The  inferomarginals  are  much  larger  and  more  prominent,  trans- 
versely oblong,  elliptical,  and  convex  or  rounded  at  the  summit,  and 
covered  with  minute,  rough,  acute  spinules,  like  those  of  the  back, 
but  rather  larger.  Of  these  about  twenty-five  to  forty  surround  the 
margin,  while  about  fifteen  to  twenty  form  a  central  row. 

A  row  of  peractinal  plates  extends  to  about  the  middle  of  the  free 
part  of  the  rays,  decreasing  rapidly  distally.  The  proximal  ones  are 


SHALLOW-WATER   STARFISHES  247 

similar  to  the  inferomarginals,  but  smaller  and  less  prominent.  The 
interradial  areas  are  rather  large,  acute-triangular,  and  covered  with 
oblong  or  elliptical  unequal  pseudopaxillae  like  the  synactinals,  but 
smaller. 

The  adambulacral  plates  have  each  two,  or  sometimes  three,  short, 
acute,  unequal,  divergent  furrow-spines,  reduced  to  one  distally. 
Their  actinal  surface  bears  a  row  of  about  six  or  seven  graded  spines, 
the  inner  one  and  that  next  to  it  being  distinctly  larger  than  the  rest 
and  standing  obliquely  on  the  plate.  These  spines  are  webbed 
together,  and  proximally  stand  in  a  curved  transverse  row. 

The  four  inner  peroral  spines  at  the  apex  of  the  jaws  are  large 
and  stout,  subacute,  the  two  median  ones  larger.  They  are  flanked 
on  each  side  by  about  six  smaller,  tapered  furrow-spines,  decreasing 
in  length  distally.  On  the  actinal  side  of  the  jaws  there  are  two  lon- 
gitudinal curved  rows  of  epioral  spines,  webbed  together,  seven  or 
eight  in  each  row,  increasing  in  size  adorally,  the  inner  one  in  each 
row  distinctly  larger. 

The  color  in  life  is  usually  either  orange-yellow  or  purple,  rarely 
lemon-yellow. 

This  species  has  a  very  extensive  geographical  distribution.  It  is 
circumpolar,  occurring  at  Greenland,  Spitzbergen,  Nova  Zembla, 
Iceland,  and  on  the  northern  coasts  of  Europe  and  Asia,  as  far 
south  as  Great  Britain  and  East  Siberia. 

On  the  Atlantic  coast  of  America  it  is  common  as  far  south  as 
Cape  Cod.  It  was  taken  by  the  U.  S.  Fish  Commission  at  numerous 
stations,  in  5  to  150  fathoms,  from  Newfoundland  to  Cape  Cod. 
Common  on  the  eastern  coasts  of  Maine  and  Massachusetts,  and  in 
the  Bay  of  Fundy,  from  low  water  to  90  fathoms,  and  off  Cape  Cod, 
20  to  50  fathoms.  Taken  on  all  the  fishing  banks  off  Nova  Scotia,  in 
20  to  150  fathoms. 

On  the  northwest  coast  of  America  it  is  common  in  Bering  Sea 
and  northern  Alaska,  and  extends  southward  to  Sitka.  I  have 
studied  specimens  from  Yakutat,  and  from  off  Juneau,  Alaska,  in  20 
fathoms  (Harriman  Expedition). 

I  have  also  examined  one  rather  large,  tumid,  nine-rayed  specimen, 
in  alcohol,  from  Bering  Island  (N.  Grebnitsky,  U.  S.  Nat.  Mus.). 
It  has  previously  been  recorded  from  Barents  Sea,  Kara  Sea,  and 
East  Siberia. 

Dr.  Fisher  records  it  from  off  the  Shumagin  Islands ;  Kasaan  Bay ; 
and  Kadiak,  Alaska,  in  12  to  123  fathoms;  and  from  Queen  Char- 
lotte Sound,  in  238  fathoms. 


248  VERRILL 

The  dorsal  surface  of  this  species  is  closely  covered  with  smaller 
pseudopaxillae,  having  finer  and  more  numerous  spinules,  than  in  any 
other  west  American  species,  except  S.  galaxides,  in  which  they  are 
still  smaller. 

It  is  allied  to  5.  stimpsoni  Verrill,  from  the  Northwest  coast  of 
America;  5".  subarcuattis  Sladen,  from  the  Southern  Indian  Ocean, 
S.  lat.  52°  04',  in  150  fathoms;  and  S.  torulatus  Sladen,  from  north 
of  the  Kermadec  Islands,  in  250  fathoms. 

TERATOLOGY. 

A  medium-sized  specimen,  taken  at  Eastport,  Maine,  has  ten  rays, 
but  one  of  the  rays  forks  beyond  the  edge  of  the  disk,  the  two  forks 
becoming  like  the  other  rays,  in  size  and  form,  distally,  so  that  it 
becomes  eleven-rayed.  Otherwise  it  has  the  ordinary  characters  of 
the  species. 

Specimens  with  nine  rays  and  twelve  rays  are  not  very  rare. 
Those  with  ten  rays  are  about  as  common  as  those  with  eleven  rays. 

SOLASTER  GALAXIDES  Verrill. 

Plate  XLVI,  figures  2,  20  (type)  ;  plate  LXXXVII,  figures  5-5<r  (details,  cotype)  ; 
plate  LXXXIX,  figure  2  (actinal  side,  type). 

Solaster  galaxides  VEWUJLL,  Amer.  Journ.  Sci.,  xxvm,  p.  59,  figs.  2,  20,  1909. 

This  is  a  broad-disked  species,  usually  with  nine  or  ten  rays,  cov- 
ered above  with  very  small,  flat-topped,  crowded  pseudopaxillae,  and 
resembling  5".  endeca  in  form  and  color.  The  two  type  specimens 
have  nine  rays.  The  larger  has  the  radii  40  mm.  and  no  mm.; 
ratio,  about  1 : 2.7.  It  was  orange-color  in  life. 

The  marginal  spines  are  about  as  in  S.  endeca,  but  the  inferomar- 
ginals  are  more  elongated  transversely,  and  bear  a  decidedly  greater 
number  of  more  minute  spinules. 

The  peractinal  series  of  pseudopaxillae  extends  only  to  about  the 
basal  third  of  the  free  part  of  the  ray.  They  are  relatively  smaller 
than  in  endeca,  being  here  only  about  half  the  size  of  the  infero- 
marginals  proximally. 

The  actinal  interradial  areas  are  apparently  relatively  larger  than 
in  endeca  of  the  same  size  and  number  of  rays,  and  bear  a  larger 
number  of  compressed  pseudopaxillae,  the  larger  ones  similar  to  the 
inferomarginals  and  peractinals.  They  form  about  sixteen  radial 
rows,  the  smaller  one  in  the  median  rows  distally.  They  are  cov- 
ered with  a  large  number  of  small,  rather  short,  regular  spinules. 


SHALLOW-WATER   STARFISHES  249 

The  dorsal  ossicles  are  unusually  small  and  closely  united,  leaving 
very  small  papular  pores,  and  the  pseudopaxillae  are  uncommonly 
small,  with  a  small  flat-topped  fascicle  of  about  twelve  to  sixteen 
minute  spinules,  of  which  three  to  six  usually  occupy  the  central  area. 

These  pseudopaxillae  are  only  about  half  as  large  as  in  typical 
Atlantic  specimens  of  endeca  of  corresponding  size.  There  are 
usually  two  subequal,  rather  long,  acute,  divergent  furrow-spines  on 
each  adambulacral  plate,  only  one  distally.  On  the  actinal  surface 
the  curved  transverse  row  or  comb  has  usually  six  to  eight  graded 
spines,  the  two  inner  decidedly  longer  and  stouter. 

The  oral  spines  and  jaw-spines  are  much  better  developed  than 
usual.  The  four  apical  spines  are  very  large,  strong,  and  acute. 
There  are  six  graded  furrow-spines  on  each  side.  The  epioral  spines 
are  long  and  slender.  They  form  two  subparallel  rows  of  about 
eight  or  nine  graded  spines.  The  spines  in  the  opposed  rows  are 
often  bent  toward  each  other  and  interlocked.  The  two  most  adoral 
are  distinctly  larger  than  the  others. 

Two  typical  specimens  from  Victoria  were  received  from  the  Pro- 
vincial Museum  of  British  Columbia.  Another,  very  similar  in 
character  and  size,  is  from  Friday  Harbor,  Puget  Sound,  cotype, 
No.  1897,  Mus.  Comp.  Zool.,  pi.  LXXXVII,  figs.  5-5C. 

SOLASTER  DAWSONI  Verrill. 

Plate  XLVI,  figures  5-56  (details)  ;  plate  xc,  figure  i  (is-rayed)  ;  plate  xci, 
figures  I,  2;  plate  xcn,  figure  i  (i3-rayed). 

Solaster  dawsoni  VERRJLL,  in  Whiteaves,  Report  Prog.  Geolog.  Survey  Canada, 
1878,  1879,  p.  4.  Whiteaves,  Trans.  Royal  Soc.  Canada,  iv,  sect.  4,  p.  1 16, 
1886.  Fisher  (pars),  op.  cit,  IQII&,  p.  313,  pi.  LXXXIV,  figs,  i,  2;  pi. 

LXXXV,  figs.  I,  2;  pi.  LXXXVI,  figs.   I,  2J  pi.  CXIII,  fig.  I. 

The  type  has  the  following  characters:  Rays  twelve.  Radius  of 
the  disk,  20  mm. ;  of  the  rays,  53  mm. ;  ratio,  about  i :  2.62.  It  is 
less  than  half  grown. 

It  has  the  general  appearance  of  S.  endeca,  as  seen  from  above, 
but  resembles  Crossaster  papposus  beneath.  The  dorsal  pseudo- 
paxillae and  their  clusters  of  stellate  spinules  are  small,  numerous, 
and  crowded.  Usually  there  are  ten  to  fifteen  minute,  short, 
webbed  spinules  around  the  edge  of  each  plate.  The  plates,  when 
denuded  of  spines,  are  small,  three-  and  four-lobed,  closely  imbri- 
cated, with  a  rounded  and  convex  central  mammilla.  The  infero- 
marginal  plates  are  prominent,  and  each  bears  a  prominent  trans- 
verse group  of  numerous  small,  slender  spinules,  forming  two  rows 
of  about  twelve  to  fourteen  each. 


250  VERRILL 

The  interradial  spaces,  beneath,  are  very  small  and  narrow,  with 
very  few  ossicles,  each  of  which  bears  a  group  of  three  or  four 
slender,  elongated  spinules.  These  spaces  are  very  much  smaller 
than  in  ^.  stimpsoni,  and  still  smaller  as  compared  with  S.  endeca. 
The  adambulacral  plates  bear  a  longitudinal  group  of  three,  or 
sometimes  four,  long,  slender,  furrow-spines,  and  outside  of  these  a 
transverse  group  of  four  or  five  rather  larger  and  longer  ones.  The 
oral  plates  bear  six  long,  slender  blunt  preoral  spines,  the  two  middle 
ones  longest,  and  four  or  five  small  lateral  ones  on  each  side. 

The  adambulacral  spines,  especially  the  furrow-series,  are  much 
longer  than  in  5".  stimpsoni  and  5".  endeca,  both  of  those  usually 
having  but  two  small  and  short  furrow-spines  on  each  plate  in  speci- 
mens of  similar  size. 

This  type  was  from  Virago  Sound,  British  Columbia,  in  8  to  15 
fathoms  (G.  M.  Dawson,  Canadian  Geological  Survey).  It  was 
dried  and  not  in  a  good  state  of  preservation.  Much  larger  and 
better  specimens  have  subsequently  been  obtained,  many  of  which 
have  been  sent  to  me  by  the  Canadian  Geological  Survey. 

One  of  the  larger  dry  specimens  from  Powell  Island,  Straits  of 
Georgia,  has  the  radii  34  mm.  and  100  mm.;  ratio,  about  i :  3.  (PI. 
xci,  fig.  2;  xcii,  fig.  i.) 

Rays  thirteen,  rather  long  and  narrow,  tapering  gradually,  the 
disk  being  smaller  and  the  rays  rather  longer  than  is  usual  in 
5".  endeca. 

The  dorsal  pseudopaxillae,  many  of  which  are  preserved  expanded, 
are  small,  crowded,  regularly  stellate,  with  a  truncate  or  infundibuli- 
form  top,  surrounded  by  about  twelve  to  eighteen  small,  slender 
spinules,  webbed  together,  and  mostly  with  one  to  three  very  small 
central  spines,  though  many  have  more. 

The  plates  are  small,  partly  four-lobed  and  partly  three-lobed, 
with  a  prominent  central  mammilla.  They  are  everywhere  closely 
imbricated,  mostly  arranged  in  quincunx  order,  and  on  the  sides  of 
the  rays  they  stand  in  close  and  pretty  regularly  alternating  oblique 
rows. 

The  papulae  are  numerous  and  small;  they  stand  partly  in  small 
groups  and  partly  singly. 

The  madreporic  plate  is  large  and  prominent,  with  numerous  fine 
gyri,  and  with  three  or  four  pseudopaxillae  on  its  surface,  near  the 
margin. 

The  inferomarginal  plates  are  large  and  very  prominent,  trans- 
versely oblong,  with  thirty  to  forty  or  more  very  slender  spinules 


SHALLOW- WATER   STARFISHES  2$  I 

around  the  margin,  and  one  or  two  smaller  rows  on  the  summit.  The 
spines  are  decidedly  longer  than  those  of  the  dorsal  surface,  espe- 
cially the  inferior  ones. 

The  superomarginal  plates  and  pseudopaxillae  are  unusually  dis- 
tinct and  well  developed.  They  are  rounded  and  bear  a  regularly 
stellate  group  of  twenty  to  thirty  slender  spinules,  longer  than  those 
of  the  dorsal  ossicles. 

The  interradial  areas  are  very  small  and  narrow,  with  a  few 
(about  twenty)  irregular  ossicles,  which  bear  round  pseudopaxillae, 
having  from  five  to  ten  long  and  slender  spinules.  None  of  the  inter- 
actinal  plates  extend  on  the  rays  beyond  the  margin  of  the  disk. 

The  furrow-series  of  adambulacral  spines  stand  mostly  four  to  a 
plate,  often  with  a  small  fifth  one  proximally ;  they  are  long,  slender, 
divergent,  webbed  together,  the  middle  ones  longest,  and  about  as  long 
as  those  of  the  transverse  series.  The  latter  are  unusually  long  and 
slender,  subequal ;  they  usually  stand  five  in  a  row,  but  often  six. 

The  six  terminal  peroral  spines  are  rather  long,  but  not  very 
stout;  they  are  regularly  graded,  and  there  are  five  or  six  in  each 
adoral  lateral  row. 

There  are  two  pairs  of  epioral  spines,  the  proximal  pair  being 
much  the  longer. 

VARIATIONS. 

A  dry,  twelve-rayed  specimen,  a  little  larger  than  the  last,  but 
having  the  same  proportions,  from  Queen  Charlotte  Islands,  has 
somewhat  larger  dorsal  pseudopaxillae,  mostly  with  twelve  to  fifteen 
somewhat  stronger  spinules.  The  furrow-series  of  spines  stand 
mostly  three  to  a  plate,  but  sometimes  four ;  they  are  about  as  long 
as,  and  rather  more  slender  than,  the  transverse  series. 

Another  specimen  from  the  same  locality  has  thirteen  rays.  The 
radii  are  28  mm.  and  85  mm. ;  ratio,  i :  3.  The  dorsal  pseudo- 
paxillae are  smaller,  elegantly  stellate,  with  about  twenty  to  twenty- 
five  slender  spinules.  The  madreporic  plate  has  a  remarkably  fine 
texture.  It  is  partly  concealed  by  a  circle  of  about  ten  stellate 
pseudopaxillae  that  stand  partly  within  its  margin.  The  furrow- 
spines  proximally  often  stand  four  on  a  plate,  though  more  com- 
monly three  to  a  plate,  the  different  rays  differing  in  this  respect  at 
corresponding  places.  The  transverse  rows  of  pectinate  spines 
contain  either  four  or  five  spines,  proximally.  The  jaws  bear  in 
some  cases  two  pairs  of  unequal  epioral  spines ;  in  others  three  pairs  ; 
or  sometimes  two  on  one  half  and  three  on  the  other  in  the  same 
individual.  The  interradial  areas  are  unusually  narrow,  and  have 


252  VERRILL 

fifteen  to  twenty  small,  round  pseudopaxillae,  with  rather  long 
spinules.  Otherwise  the  characters  are  as  in  the  type. 

A  dry,  thirteen-rayed  specimen,  in  poor  condition,  from  Esqui- 
mault  Harbor,  has  unusually  small  dorsal  pseudopaxillae,  so  crowded 
that  when  the  regularly  stellate  spinules  are  expanded  the  adjacent 
stars  touch  each  other  by  their  edges,  or  interlock,  over  most  of  the 
surface. 

A  fifteen-rayed  young  specimen  (radii,  47  mm.  and  18  mm.)  from 
Vancouver  Island  (see  fig.)  has  four  of  the  contiguous  rays  on  one 
side  shorter  than  the  rest,  the  two  intermediate  equal  ones  shorter 
than  the  next  on  each  side,  which  are  also  nearly  equal,  thus  giving 
the  specimen  a  bilateral  symmetry.  The  oral  spines  are  four  on  each 
jaw,  rather  stout,  nearly  equal;  epiorals  two  to  four,  also  rather 
stout ;  laterals  small,  slender,  graded,  about  six  on  each  side.  Actinal 
interradial  pseudopaxillae  are  few  and  very  small.  Adambulacral 
groove-spines  about  three  in  middle  of  rays,  rather  strong,  often 
four  proximally ;  comb-spines  about  four,  subequal.  Dorsal  pseudo- 
paxillae very  numerous,  small,  nearly  even,  circular ;  those  on  the  rays 
very  small  and  crowded  distally. 

A  somewhat  smaller  thirteen-rayed  specimen  (radii,  42  mm.  and 
16  mm.),  from  the  same  locality,  agrees  well  with  the  last  in  the 
dorsal  and  actinal  paxillae,  but  has  relatively  smaller  and  more 
slender  adambulacral  and  oral  spines.  The  apical  jaw-spines  are 
mostly  six  on  each  jaw,  small  and  slender. 

This  species  has  a  wide  range,  especially  from  Monterey  Bay, 
California,  to  the  Aleutian  Islands.  It  is  common  in  the  waters  of 
Puget  Sound  and  British  Columbia.  I  have  studied  specimens,  sent 
by  the  Canadian  Geological  Survey,  from  Virago  Sound;  Queen 
Charlotte  Islands;  Powell  Island,  Gulf  of  Georgia  (G.  M.  Dawson). 
Esquimault  Bay  (C.  F.  Newcombe).  Vancouver  Island;  Departure 
Bay  (Geological  Survey  of  Canada)  ;  and  from  other  localities. 

Whiteaves  (1886)  recorded  it  from  Powell  Island,  Malaspina 
Inlet,  and  Galatos  Channel  (abundant  at  low  tide). 

Fisher  records  it  from  many  localities  from  Monterey  Bay  to  the 
Aleutian  Islands,  from  low  tide  to  123  fathoms.  He  also  records  it 
from  more  northern  localities;  but  some,  if  not  most,  of  these 
localities  refer  to  S.  dawsoni  arctica  Verrill,  perhaps  a  distinct 
species,  which  he  did  not  distinguish. 

His  more  northern  localities  include  the  Kuril  Islands  (229 
fathoms)  ;  Shumagins ;  and  Point  Franklin,  on  the  Arctic  Ocean. 
The  latter  is  the  type-locality  of  S.  danvsoni  arctica  Verrill. 


SHALLOW-WATER   STARFISHES 


253 


SOLASTER  DAWSONI  ARCTICA  Verrill,  subsp.  nov. 
Plate  LXXXVII,  figures  6,  6a  (details)  ;  text-figure  No.  14. 

This  subspecies  is  very  similar  in  general  appearance  to  5".  endeca 
and  S.  dawsoni.  Its  dorsal  pseudopaxillae  are  rather  larger  than  in 
the  former  and  are  flat-topped.  The  larger  dorsal  ones,  on  the  disk 
and  bases  of  the  rays,  are  covered  with  twenty  to  thirty,  or  more, 
small,  short,  very  slender,  thorny  spinules,  of  which  about  fifteen  to 
twenty-five  are  marginal,  the  rest  central;  all  are  short,  rising  to 
about  the  same  height,  but  the  central  ones  may  be  a  little  shorter, 
in  some  cases.  Papulae  are  small  and  stand  singly. 

The  adambulacral  plates  bear  two  rather  long  and  slender,  nearly 
equal,  furrow-spines,  sometimes  with  a  smaller  supplementary  one, 


FIG.  14. 

Solaster  dawsoni  arctica  Ver.,  type,  Arctic   O.     U.   S.   Nat.   Mus.     Two  adambulacral 
plates  and  their  spines;  a,  a,  outer  or  actinal  combs;  b,  b',  inner  or  furrow  groups.     X  17. 

and  a  transverse  comb  of  about  five  or  six  somewhat  shorter  and 
stouter  spines,  not  very  diverse  in  length,  so  that  their  tips  lie  in  a 
slightly  convex  line.  The  furrow-spines  rise  to  about  the  same 
height. 

The  inferomarginals  are  relatively  rather  large  and  prominent, 
strongly  compressed,  transversely  oblong,  covered  with  a  great  num- 
ber of  minute,  rough  spinules,  in  three  or  four  transverse  rows.  The 
superomarginals  are  small,  but  distinct,  not  half  as  large  as  the  lower 
ones,  but  twice  as  large  as  the  dorso-laterals,  spinulated  like  the 
lower  ones. 

Peractinals  are  small  and  inconspicuous,  except  proximally.  The 
oral  plates  have  about  six  or  seven  graded  spines  on  each  end,  besides 


254  VERRILL 

four  stout  apical  ones.  Epioral  spines  numerous,  six  to  eight  in 
each  cluster. 

Radii  of  the  type  are  20  mm,  and  55  mm. ;  ratio,  2 :  2.75 ;  rays  ten. 

Northern  Alaska.  The  type  is  from  near  Point  Franklin,  Arctic 
Ocean,  in  13^  fathoms  (coll.  Murdoch,  Point  Barrow  Expedition, 
U.  S.  Nat.  Mus.,  No.  7624). 

This  form,  which  may  prove  to  be  a  distinct  species,  is  easily  dis- 
tinguished from  typical  6".  dawsoni  by  the  much  finer  and  more 
numerous  spinules  of  the  dorsal  pseudopaxillse.  They  are  only  about 
half  as  large,  but  nearly  twice  as  numerous.  The  less  projecting 
adambulacral  combs,  the  general  presence  of  only  two  nearly  equal 
furrow-spines,  and  other  less  notable  characters  also  serve  to  dis- 
tinguish it. 

Professor  Fisher  records  a  specimen,  from  the  same  locality,  as 
5*.  dawsoni,  which  may  be  identical  with  this.  The  locality  is  far 
more  northern  and  arctic  than  any  known  for  the  true  dawsoni. 
My  material  is  not  sufficient  for  a  positive  decision  as  to  its  specific 
distinctness. 

It  is  certainly  more  closely  related  to  S.  dawsoni  than  to  S.  endeca, 
though  the  latter  is  the  common  species  of  the  arctic  coasts,  while 
S.  dawsoni  is  common  from  the  Aleutian  Islands  southward  to  Cali- 
fornia. 

SOLASTER  STIMPSONI  Verrill. 

Plate  x,  figures  i,  2;  plate  xi,  figures  I,  2;  plate  xv,  figures  I,  2;  plate  XLVI, 
figures  i-ic  (details)  ;  plate  xov,  figure  2  (type)  ;  plate  xcv  (type). 

f  Asterias  decemradiatus  BRANDT,  Prod.,  p.  271,  1835  (no  description). 
Solaster  stimpsoni  VERRILL,  in  Whiteaves,  Invert,   of  Queen   Charlotte  Is., 

Rep.  Prog.  Geol.  Survey  Canada,  for  1878-79,  p.  3;  Whiteaves,  Trans. 

Royal  Soc.  Canada,  iv,  p4  116,  1887.     Fisher,  op.  cit,  ignb,  p.  311,  pi. 

LXXXII,  fig.  3;  pi.  LXXXIII,  figs.  1-5. 
f  Solaster  decemradiatus  STIMPSON,  Journ.  Boston  Soc.  Nat  Hist.,  vi,  p.  89, 

1857   (no  description). 
Solaster  vancouverensis  DE  LORIOL,  Mem.  Soc.  Phys.  Geneve,  xxxn,  part  2, 

No.  9  [p.  12],  pi.  i,  [xvi],  fig.  5,  1896. 

Disk  of  moderate  width;  rays  elongated,  usually  ten,  sometimes 
nine.  One  of  the  type  specimens  has  the  radii  of  the  disk  30  mm. ; 
of  the  different  rays,  100  mm.  to  115  mm.;  ratios,  usually  about 

i :  3-7S-4- 

The  rays  are  long,  rounded  above,  regularly  tapered,  upper  sur- 
face thickly  covered  with  pseudopaxillae  composed  of  stellate  clusters 
of  small,  blunt,  strongly  divergent,  webbed  spinules,  the  larger  ones, 
on  the  rays,  having  usually  five  to  eight  spinules  around  the  edge 


SHALLOW-WATER   STARFISHES  255 

and  one  or  two  in  the  middle  of  the  star ;  on  the  disk,  where  they 
are  more  crowded,  they  may  have  six  to  ten  radial  spinules,  with  two 
or  three  central  ones.  In  the  smaller  specimens  there  are  fewer 
spinules  in  the  clusters.  These  spinules  are  decidedly  larger,  stouter, 
more  obtuse  and  less  numerous  than  in  S.  endeca,  and  form  larger 
pseudopaxillae.  On  the  lower  side,  the  interradial  spaces  are  smaller 
than  in  5".  endeca,  with  fewer  and  smaller  ossicles,  which  are 
roundish.  Each  bears  a  stellate  group,  usually  with  four  or  five  taper- 
ing spinules  around  the  edge  and  one  or  two  central  ones.  They  are 
rather  larger  than  those  of  the  dorsal  surface,  and  much  stouter  and 
fewer  than  the  corresponding  spinules  of  S.  endeca. 

The  inferomarginal  plates  are  smaller  and  less  prominent  than  in 
S.  endeca.  Each  bears  a  transverse  group  of  about  twelve  to  sixteen 
spinules,  mostly  in  two  rows,  similar  to  those  of  the  sides  and  under 
surface  of  the  rays. 

Each  adambulacral  plate  bears,  on  its  inner  edge,  two  small,  short, 
tapered,  acute  spines,  rarely  three  proximally,  which  form  a  longi- 
tudinal row ;  and  on  the  actinal  surface  of  each,  a  simple  transverse 
row  of  about  six  to  eight  longer  and  larger,  subequal,  tapered  spines, 
with  subacute  tips,  the  inner  one  a  little  larger.  These  are  shorter, 
less  unequal,  stouter  and  less  acute  than  those  of  S.  endeca.  The 
jaws  bear  six  strong  and  rather  long  peroral  spines,  the  two  middle 
ones  longest,  the  others  graded ;  and  about  six  smaller  graded  spines 
on  each  side. 

VARIATIONS. 

The  largest  specimen  that  I  have  seen  is  from  Victoria  (coll.  C.  F. 
Newcombe,  Prov.  Mus.  B.  C.).  Its  radii  are  42  mm.  to  45  mm.  and 
180  mm.  to  190  mm. ;  ratios,  about  i :  4.25. 

The  ten  rays  are  long,  the  distal  part  slender.  It  has  been  stained 
(to  imitate  the  natural  colors)  bright  orange,  with  a  central  star  and 
a  median  band  on  each  ray  of  dark  purple.  The  dorsal,  stellate 
pseudopaxillae  are  larger  than  in  the  type,  and  not  so  closely 
crowded.  The  larger  ones  (pi.  XLVI,  fig.  ia)  mostly  bear  five 
divergent  webbed  spinules  around  the  edge  and  one  in  the  middle, 
making  a  rather  large,  but  short,  regularly  star-shaped  group. 
They  stand  in  pretty  regular  obliquely  transverse,  alternating  rows 
on  the  sides  of  the  rays,  and  in  longitudinal  rows  on  the  dorsal  side ; 
also  in  regular  quincunx  order  in  many  places  on  the  sides.  The 
papulae  mostly  stand  singly,  but  often  two  or  three  stand  together, 
especially  on  the  disk. 


256  VERRILL 

The  madreporic  plate  is  large  and  has  four  pseudopaxillse  on  its 
surface,  and  others  irregularly  placed  at  its  margin. 

The  adambulacral  spines  in  the  furrow-series  are  short,  and  stand 
mostly  three  to  a  plate,  the  middle  one  longest;  but  at  about  the 
distal  fourth  of  the  rays  they  are  reduced  to  two.  The  actinal  trans- 
verse combs  consist  proximally  of  about  eight  to  ten  webbed  and 
feebly  graded  spines,  in  a  simple,  slightly  curved  row,  the  inner 
ones  slightly  longer  and  larger  than  the  middle  ones ;  but,  owing  to 
the  convex  summit  of  the  plate,  the  middle  spines  are  often  more 
prominent  and  the  tips  form  a  curved  outline;  the  outer  ones  are 
much  shorter. 

The  inferomarginal  plates  are  transversely  oblong  and  often  com- 
pressed. Each  bears  a  marginal,  webbed  series  of  about  eight  to  ten 
rather  strong  spinules  on  each  side,  and  often  a  partial  median  row 
of  two  to  four  spinules. 

The  superomarginal  plates  are  small,  rounded  or  ovate,  close  to  the 
inferomarginals,  and  bear  a  stellate  group  usually  of  five  to  seven 
spinules,  like  those  of  the  dorsal  surface. 

The  actinal  interradial  areas  are  relatively  small  and  narrow,  and 
are  covered  by  small,  rounded,  unequal  pseudopaxillae,  which  form 
about  eight  radial  rows  distally.  They  mostly  bear  four  to  six, 
rarely  eight,  stout  spinules  in  a  stellate,  webbed  group,  rarely  with  a 
central  one.  Their  spinules  are  like  those  of  the  adjacent  infero- 
marginal plates,  but  fewer. 

The  peroral  spines  consist  of  a  terminal  group  of  six  stout,  close, 
graded,  subacute  spines,  with  three  or  four  much  smaller  spines  on 
each  side.  The  epiorals  consist  of  two  opposed  curved  and  con- 
vergent rows  of  seven  to  nine  graded  spines,  often  with  the  opposed 
tips  in  contact  or  interlocking,  the  two  inner  ones  decidedly  stouter. 

Pedicellariae  of  minute  size  occur  in  this  species.  They  were  first 
observed  by  A.  H.  Verrill,  while  drawing  the  details  of  the  dorsal 
pseudopaxillae  mounted  for  the  microscope.  They  are  very  small, 
sessile  around  the  papular  pores,  bivalve,  often  with  the  valves 
unequal.  (See  pi.  XLVI,  fig.  ic.)  They  a^e  scarcely  visible  to  the 
naked  eye  and  occur  only  in  small  numbers,  so  far  as  observed,  and 
are,  therefore,  likely  to  be  overlooked.  Some  specimens  seem  to  lack 
them  entirely. 

This  species  has  a  wide  range,  from  Oregon  to  the  Aleutian 
Islands,  and,  according  to  Fisher,  has  been  taken  also  farther  north, 
at  Bering  Island,  Commander  Islands  group. 

Dr.  Fisher  records  it  from  low  tide  to  33  fathoms,  and  from 
Oregon  to  Bering  Island. 


SHALLOW-WATER   STARFISHES  257 

It  is  especially  characteristic  of  the  region  from  Puget  Sound  to 
southern  Alaska,  at  low  tide  and  in  shallow  water.  I  have  received 
numerous  good  specimens  from  the  Canadian  Geological  Survey, 
taken  at  Departure  Bay,  Vancouver  Island,  etc.  Also,  from  C.  F. 
Newcombe,  specimens  from  off  Victoria,  Vancouver  Island  (large) . 
The  type  was  from  Ramsay  Island,  British  Columbia  (Canadian 
Geological  Survey). 

SOLASTER  CONSTELLATUS  Verrill. 

Plate  XLVI,  figures  3,  4  (type)  ;  plate  xc,  figure  2  (type)  ;  plate  xan  (type) ; 
plate  xciv,  figure  I   (type). 

Solaster  constellates  VERRILL,  Amer.  Journ.  Sci.,  xxvni,  p.  60,  figs.  3,  4,  1909. 
Fisher,  op.  cit,  191  ib,  p.  311,  foot-note. 

The  type  is  eight-rayed,  with  a  small  disk  and  long,  tapered  arms. 
Radii  of  the  type  are  21  mm.  and  78  mm. ;  ratio,  1 : 3.7. 

The  dorsal  pseudopaxillae  are  unusually  large  and  elevated,  regular 
and  flower-like.  They  are  stellate  in  form,  and  usually,  where 
largest  on  the  disk  and  base  of  rays,  they  have  a  single  central 
spinule  and  about  six  equally  spaced  and  webbed  marginal  spinules, 
which  are  often  fully  expanded  and  nearly  horizontal,  producing 
the  appearance  of  a  six-petaled  flower.  The  largest  ones  may  have 
six  to  eight  divergent  spines,  and  the  small  distal  ones  only  four  or 
five.  The  superomarginal  and  interactinal  ones  are  quite  similar  to 
the  dorsals.  The  peractinals  are  small. 

The  dorsal  papulae  are  small  and  mostly  stout,  and  stand  singly 
around  the  pseudopaxillae.  The  latter  are  very  regularly  arranged  in 
oblique  alternating  rows  on  the  sides  of  the  rays,  or  in  quincunx. 

The  inferomarginal  plates  are  small,  roundish,  and  bear  a  small 
number  (eight  to  twelve)  of  elongated,  webbed  spinules  similar  to 
the  dorsal  ones,  but  longer. 

The  adambulacral  spines  consist  of  a  furrow-series  with  two,  or 
sometimes  three,  short,  tapered  spines,  and  an  outer  comb  of  six  to 
eight  nearly  equal,  tapered  spines,  webbed  nearly  to  the  tips;  the 
inner  ones  are  usually  rather  longer,  so  that  the  rows  are  a  little 
graded.  Adoral  spines  are  strongly  graded,  about  ten  to  a  jaw,  the 
apical  ones  unusually  stout.  The  superomarginal  pseudopaxillae  are 
but  little  larger  than  the  dorsal  ones,  but  are  distinct,  partly  owing 
to  their  greater  elevation. 

This  is  not  the  only  eight-rayed  species  known  to  me  from  that 
coast.  Its  large  and  beautifully  stellate  dorsal  pseudopaxillae  are 
distinctive  and  decidedly  larger  and  with  longer  spinules  than  in 
18 


258  VERRILL 

S.  stimpsoni  Verrill,  which  it  somewhat  resembles  in  certain  respects. 
The  latter  is  also  sometimes  eight-rayed. 

The  type  is  from  Puget  Sound  (Prof.  T.  Kincaid,  University  of 
Washington). 

Fisher  (1911  b,  p.  311)  suggests  that  this  is  merely  an  individual 
eight-rayed  variation  of  S.  stimpsoni.  The  species  was  not  based 
on  the  eight-rayed  condition,  for  no  one  knows  better  than  the  writer 
the  variability  of  species  of  Solaster  in  that  respect.  Perhaps  no  one 
has  collected  and  studied  more  specimens  of  the  genus.  This  may 
be  but  a  variety  of  stimpsoni,  but  among  large  numbers  of  the  latter 
studied,  from  the  same  region,  no  intermediate  specimens  have  been 
found.  The  most  notable  difference  is  in  the  larger  size  of  the 
stellate  pseudopaxillae,  with  their  few,  long,  slender,  divergent, 
webbed  spinules. 

SOLASTER  PAXILLATUS  Sladen. 

Solaster  paxillatus  SLADEN,  Voy.  Challenger,  xxx,  p.  452,  pi.  LXXI,  figs.  1-3; 
pi.  LXXII,  figs,  i,  2,  1889.  Fisher,  op.  cit.,  19116,  p.  315,  pi.  LXXXVII,  figs. 
I,  2;  pi.  LXXXVIII;  pi.  LXXXIX,  figs.  I,  2;  pL  cxm,  fig.  3. 

Rays  eight  to  ten.  Disk  of  rather  large  size.  Radii,  58  mm.  and 
117  mm. ;  ratios,  about  1 : 3,  sometimes  up  to  1 :  3.7. 

Dorsal  pseudopaxillae  are  numerous,  rather  close,  subtabulate,  a 
little  convex ;  their  spinules  are  numerous  and  short,  thickly  webbed 
together,  about  thirty  to  forty  on  the  larger  plates. 

Marginal  plates  prominent,  alternate,  very  unequal  in  size,  upper 
ones  about  as  large  as  dorsal  paxillae.  Inferomarginals  compressed, 
fan-shaped,  with  thirty  to  forty  spinules.  They  form  a  prominent 
border  to  the  rays. 

Adambulacral  plates  with  three  or  four  long  spines  in  the  furrow- 
series.  Actinal  series  in  fans  of  five  to  eight  proximally,  long, 
tapered,  acute,  standing  in  a  curved  line,  the  inner  end  turned 
aborally. 

Dr.  Fisher  records  this  species  from  seven  stations,  from  Bering 
Sea  to  Kadiak,  Alaska,  in  5  to  276  fathoms.  The  type  was  from  off 
Yokohama,  Japan,  in  344  fathoms. 

The  above  description  is  condensed  from  Fisher. 

Genus  Crossaster  (Muller  and  Troschel)  restr.  Verrill. 

Crossaster  (pars)  MULLER  and  TROSCHEL,  op.  cit.,  18400,  p.  103. 

Solaster  (pars)  FORBES,  Mem.  Wern.  Soc.,  vni,  p.  129,  1839;  Brit.  Starfishes, 

p.  112,  1841.    Muller  and  Troschel,  Syst.  Aster.,  p.  26,  127,  pi.  in,  figs. 

I,  a-b,  1842.  Fisher,  op.  cit.,  19116,  pp.  306,  320. 


SHALLOW-WATER   STARFISHES  259 

Crossaster  VERRILL,  Proc.  Boston  Soc.,  x,  pp.  345,  356,  1866  (here  first  re- 
stricted). A.  Agassiz,  N.  American  Starfishes,  p.  98,  pi.  xn  (structure), 
1877.  Duncan  and  Sladen,  op.  cit.,  p.  36,  1881.  Sladen,  Voy.  Challenger, 
xxx,  p.  444,  1889. 

Solaster  (pars)  DANIELSSEN  and  KOREN,  p.  48,  pi.  ix,  1884. 

Rays  variable  in  number,  usually  ten  to  twelve.  Dorsal  skeleton 
feebly  developed,  flexible,  the  ossicles  slender  and  openly  reticulated, 
leaving  numerous  rather  large  papular  areas.  The  dorsal  and  mar- 
ginal plates  bear  elongated  pseudopaxillae,  with  numerous  slender 
spinules  clustered  in  the  form  of  a  brush  or  pencil,  the  middle  ones 
longest.  Superomarginals  feebly  developed;  inferomarginals  much 
larger,  with  large  pencils  of  spinules;  actinal  plates  few,  sometimes 
lacking.  Adambulacral  spines  form  a  regular  furrow-series,  usually 
three  to  five  to  a  plate,  and  an  exterior  transverse  comb  of  longer, 
webbed  spines. 

The  dorsal  ossicles  do  not  form  definite,  oblique,  transverse  rows 
on  the  sides  of  the  rays,  as  they  do  in  Solaster, 

CROSSASTER  PAPPOSUS  (Linne)  Miiller  and  Troschel. 
Plate  v,  figure  2 ;  plate  vm,  figures  i,  2 ;  plate  ix,  figure  4 ;  plate  XLIX,  figure  4. 

Asterias  papposa  LINNE,  Syst.  Nat.,  ed.  xn,  p.  1098,  1767.    Walch,  p.  76,  1774. 

0.  F.  Miiller,  op.  cit.,  p.  234,  1776.    Fabricius,  Fauna  Groenlandica,  p.  369, 
1780. 

Solaster  papposus  FORBES,  Mem.  Wern.  Soc.,  vm,  p.  121,  1839;  Brit.  Starfishes, 
p.  112,  fig.,  1841.  Stimpson,  Invert.  Grand  Manan,  p.  15,  1853.  Muller 
and  Troschel,  Syst.,  p.  26,  1842.  Ltitken,  Oversigt  over  Gronlands  Echino- 
dermata,  p.  40,  1857.  Perrier,  Stellerides  du  Mus.,  Arch.  Zool.  Exper., 
p.  94.  Viguier,  Squelette  des  Stell.,  op.  cit.,  p.  124,  1878.  Danielssen  and 
Koren,  op.  cit.,  p.  48,  pi.  ix,  fig.  12,  1884.  Fisher,  op.  cit,  191  ib,  p.  325, 
pi.  xciv,  figs.  1-6. 

Solaster  (Polyaster)  papposus  GRAY,  op.  cit.,  p.  183,  1840;  Synopsis,  p.  5,  1866. 

Crossaster  papposus  MULLER  and  TROSCHEL,  Wieg.  Arch.,  iv,  part  i,  p.  183, 
1840.  Verrill,  Proc.  Boston  Soc.  Nat.  Hist,  x,  p.  345,  1866.  A.  Agassiz, 
North  Amer.  Starfishes,  pp.  99,  112,  pi.  xn,  figs.  1-5,  1877.  Bush,  op.  cit, 
p.  246,  1883.  Verrill,  Expl.  Albatross,  Rep.  U.  S.  Fish  Comm.  for  1883, 
p.  541,  1885.  Murdoch,  op.  cit,  p.  159,  1885.  Fewkes,  op.  cit.,  p.  63,  fig., 
1891.  Duncan  and  Sladen,  op.  cit,  p.  36,  pi.  in,  figs.  1-4,  1881.  Sladen, 
Voy.  Challenger,  Zool.,  xxx,  p.  444,  1889.  Ganong,  Echinod.  New  Bruns- 
wick, p.  31,  fig.,  1898.  Verrill,  Amer.  Journ.  Sci.,  XLIX,  p.  201,  1895  (dis- 
tribution). Perrier,  op.  cit,  p.  40,  1896. 

Crossaster  papposus  DODERLEIN,  op.  cit.,  p.  338,  1899.    Ludwig,  Fauna  Arctica, 

1,  p.  460,  1900  (synonymy  and  distribution). 

Asterias  aftnis  and  f  Asterias  alboverrucosa  BRANDT,  Prod.  Descr.  Anim., 
p.  271,  1835  (young,  described  from  colored  figures). 


200  VERRILL 

A  typical  Atlantic  specimen,  of  medium  size,  has  the  radii  30  mm. 
and  55  mm.,  but  the  proportions  vary  considerably.  Rays  thirteen, 
tapered,  acute.  The  whole  upper  surface  is  covered  with  rather 
large,  unequal,  elevated,  spaced  pencils  or  conical  tufts  of  slender, 
elongated  spinules,  more  or  less  divergent,  pointed,  forming  rather 
high  pseudopaxillae.  Superomarginal  plates  small,  not  easily  dis- 
tinguishable; lower  ones  larger,  well  defined,  spaced,  bearing  large 
tufts  of  spinules.  Actinal  spinules  mostly  lacking.  Furrow-series 
of  adambulacral  spines  consists  of  four  or  five  slender  subequal 
spines  on  each  plate.  The  transverse  comb  has  six  or  seven  longer, 
slender,  acute  spines.  The  four  apical  adoral  spines  are  long  and 
pretty  stout;  they  are  flanked  on  each  side  by  about  six  slender 
graded  spines. 

In  life  this  is  a  very  beautiful  species.  The  color  in  life  is  usually 
light  red  or  pale  orange,  with  the  tips  of  the  rays  bright  red ;  under 
side,  light  yellow.  It  sometimes  becomes  nearly  a  foot  in  diameter. 

The  Pacific  specimen  figured  (pi.  vui,  figs,  i,  2;  pi.  ix,  fig.  4)  was 
taken  in  Berg  Bay,  at  10  fathoms,  June  10,  1899  (W.  R.  Coe,  Harri- 
man  Expedition).  According  to  Dr.  Coe's  notes  it  was,  in  life: 
"  Pale  yellow,  with  a  horseshoe-shaped  red  spot  on  the  middle  of  the 
disk,  and  several  small  pale  pink  spots  on  the  rays." 

Its  radii  are  16  mm.  and  30  mm.,  in  alcohol.  In  all  essential 
characters  it  agrees  well  with  the  Atlantic  specimens  of  similar  size. 
It  has  the  tufts  of  spinules  of  the  pseudopaxillae  on  the  dorsal  surface 
expanded  and  mostly  divergent,  so  that  the  tufts  appear  broader  and 
more  stellate  than  in  dry  specimens.  The  papulae  are  small,  rather 
numerous,  mostly  standing  singly,  or  two  together,  on  the  wide  papu- 
lar areas,  around  the  pseudopaxillae. 

This  species  varies  much  in  appearance,  according  to  the  mode  of 
preservation.  Owing  to  the  feeble  skeleton,  it  is  apt  to  become 
flaccid  and  soft  in  drying,  while  the  dorsal  spinules  may  droop  to  one 
side,  or  collapse,  so  as  to  give  very  unnatural  appearances.  The 
number  of  rays  varies  from  nine  to  fourteen,  but  they  are  usually  ten 
to  twelve. 

This  species  is  circumpolar  and  has  a  very  extensive  geographical 
range.  It  extends  southward  on  the  European  coasts  to  Scandinavia, 
Ireland,  Great  Britain,  and  France.  It  is  known  from  Greenland, 
Iceland,  Spitzbergen,  Barents  Sea,  East  Siberia,  etc.  It  extends 
southward  to  Cape  Cod,  and  beyond,  in  deep  water,  on  the  American 
side.  It  is  not  uncommon  in  the  Bay  of  Fundy  on  the  shore,  on  stony 
or  nullipore  bottoms,  at  very  low  tides,  where  I  have  often  taken 


SHALLOW- WATER   STARFISHES  26 1 

fine  specimens;  but  it  is  more  common  in  10  to  50  fathoms  or  more. 
Large  specimens  are  often  taken  on  the  Newfoundland  Banks  and 
the  fishing  banks,  off  Nova  Scotia,  by  the  fishermen,  in  40  to  100 
fathoms,  on  their  deep-sea  lines.  It  was  taken  by  the  U.  S.  Fish 
Commission,  at  many  stations,  in  5  to  179  fathoms,  mostly  on  hard 
bottoms,  from  Newfoundland  Banks  to  off  Cape  Cod. 

On  the  Pacific  side  it  occurs  from  the  Arctic  Ocean  south  to 
Vancouver  Island,  in  shallow  water;  and  on  the  coast  of  Siberia. 

I  have  examined  specimens  from  many  localities:  Bering  Island, 
eleven-rayed  (N.  Grebnitsky) ;  Point  Franklin  (Murdoch),  No. 
7626;  and  Cape  Prince  of  Wales  (W.  H.  Ball),  No.  6059,  U.  S. 
National  Museum ;  near  Victoria,  British  Columbia,  one,  nine-rayed ; 
one,  ten-rayed;  one,  eleven-rayed  (Prov.  Mus.  B.  C.)  ;  Berg  Bay, 
eleven-rayed  (W.  R.  Coe)  ;  Puget  Sound  (Mus.  Comp.  Zool.)  ; 
several,  good;  Vancouver  Island  (Professor  J.  Macoun,  Canadian 
Geological  Survey,  1909),  many  good. 

Dr.  Fisher  records  it  from  certain  localities,  from  Bering  Strait 
to  Washington,  and  from  low  tide  to  283  fathoms;  but  mostly  in 
from  5  to  50  fathoms,  on  hard  bottoms. 

Asterias  affinis  and  A.  alboverrucosa  Brandt  were  probably 
identical  with  this  species  of  Crossaster,  but  the  descriptions  were 
very  brief  and  imperfect,  having  been  based  entirely  on  figures  of 
young  specimens.  Both  were  one  inch  in  diameter  of  disk ;  length  of 
rays,  seven  lines.  Both  were  described  as  having  ten  rays,  with 
large,  scattered  clusters  of  dorsal  spinules  ("papillae"),  and  as 
resembling  A.  papposa.  The  number  of  rays  cannot  be  regarded  as 
a  specific  character.  From  the  descriptions,  both  might,  without 
hesitation,  be  referred  to  C.  papposus,  which  is  the  form  found  in 
the  same  region,  Bering  Sea,  where  his  supposed  species  were  found, 
and  where  no  other  similar  species  has  been  obtained  by  modern  col- 
lectors. 

The  Crossaster  affinis  Danielssen  and  Koren,1  from  the  North 
Atlantic  (C.  koreni  Verrill),  is  not  to  be  considered  the  same  as 
Brandt's  a/finis,  which  appears  to  be  an  ordinary  ten-rayed  papposus, 
the  difference  in  the  number  of  rays  being  the  only  difference  given, 

1  This  is  referred  to  C.  papposus  by  Ludwig  and  others,  but  the  differences, 
as  described  and  figured  by  Danielssen  and  Koren  (1884,  pp.  44-47,  pi.  ix, 
figs.  7,  8,  14),  seem  to  be  of  specific  value — notably  the  shorter  and  closer 
dorsal  paxillae;  the  increased  number  and  shorter  adambulacral  spines;  the 
larger  actinal  interradial  areas,  with  larger  ossicles ;  and  the  trifid  odontophore. 

As  it  seems  to  have  had  no  tenable  name,  I  would  propose  for  it  C.  KORENI. 
It  has  not  been  found  on  the  Atlantic  coast  of  America. 


262  VERRILL 

and  this  is  also  a  common  number  of  rays  in  the  common  Atlantic 
papposus. 

Ludwig  (Fauna  Arctica,  p.  464)  identified  Brandt's  alboverru- 
cosa  with  S.  endeca;  but  Brandt  recorded  the  latter  (var.  decem- 
radiata)  as  from  Sitka,  in  the  same  article.  His  descriptions  are, 
of  themselves,  wholly  insufficient  to  identify  any  species,  and  prob- 
ably the  colored  drawings  on  which  they  were  based  were  neither 
accurate  nor  detailed.  It  is  possible  that  the  names  on  the  original 
drawings  have  been  transposed  since  Brandt  named  them  in  1835. 

A  closely  related  species  (S.  penicillatus  Sladen)  occurs  in  the 
South  Atlantic,  S.  lat.  37°  25'  30"  to  46°  43',  in  no  to  140  fathoms. 

Family  ASTERINIDJE  Gray. 

Asterinida  GRAY,  Ann.  and  Mag.  N.  Hist.,  vi,  p.  228,  1840;  Synopsis,  p.  15, 1866. 
Perrier,  Revis.  Stell.,  iv,  p.  292,  1875;  v,  p.  209,  1876.  Viguier,  Squelette 
des  Stell.,  Arch.  Zool.  Exper.,  vn,  p.  205,  pi.  xiv,  figs.  1-13,  1878  (structure). 
Sladen,  Voy.  Challenger,  xxx,  p.  374,  1889.  Perrier,  Expl.  Trav.  et 
Talism.,  pp.  141,  163,  1894.  Fisher,  op.  cit.,  19116,  p.  253  (table  of  genera). 
Verrill,  Revision  of  the  Genera,  Amer.  Journ.  Science,  xxxv,  p.  477,  May, 
1813. 

Body  usually  rather  flat,  often  thin,  normally  pentagonal  or  hex- 
agonal, rarely  with  eight  rays ;  edges  usually  thin,  formed  mainly  by 
the  inferomarginals ;  marginal  plates  small,  usually  scarcely  larger 
than  the  adjacent  dorsals,  covered  with  a  group  of  spinules.  Dor- 
sal plates  usually  flat  and  more  or  less  imbricated,  sometimes  not 
imbricated ;  generally  covered  with  granules  or  minute  spinules,  not 
rarely  with  tufts  or  combs  of  spines ;  sometimes  covered  with  a  soft, 
naked,  or  granular  dermis.  Under  side  flat;  actinal  plates  imbricated 
or  closely  united,  usually  furnished  with  small  combs  or  tufts  of 
spinules,  or  with  only  one.  Adambulacral  spines  generally  webbed ; 
furrow-spines  form  a  comb  of  two  to  six  or  more,  sometimes  two  or 
more  combs  superimposed.  Pedicellariae  usually  lacking;  when 
present  erect,  two-bladed. 

Genus  Asterina  Nardo. 
Type,  A.  minuta  Nardo  =  A.  gibbosa  (Penn.). 

Asterina  NARDO,  Oken's  Isis,  p.  716,  1834.  Gray,  Ann.  and  Mag.  Nat.  Hist., 
vi,  p.  286,  1840;  Synopsis,  p.  16,  1866.  Perrier,  Revis.  Stell.,  Arch.  Zool. 
Exper.  et  Gen.,  v,  p.  214.  1876.  A.  Agassiz,  North  American  Starfishes, 
p.  106,  pi.  xiv,  1877  (structure).  Viguier,  op.  cit.,  vn,  p.  207,  pi.  xiv, 
figs.  8-13,  1878  (structure).  Sladen  (pars),  Voy.  Challenger,  xxx,  p. 
388,  1889.  Fisher  (pars),  op.  cit.,  19116,  p.  254. 


SHALLOW- WATER   STARFISHES  263 

Ctenaster  L.  AGASSIZ,  Prod.,  p.  192,  1835  (type,  C.  mintitus). 

Asteriscus  (pars)  MULLER  and  TROSCHEL,  1840,  p.  104;  Syst.  Aster.,  p.  39,  1842. 

Asterina  (restricted)  VERRILL,  Amer.  Journ.  Science,  xxxv,  May,  1913,  p.  481. 

The  genus  Asterina  should  be  restricted  to  those  species  closely 
allied  to  A.  gibbosa  (Penn.)  of  southern  Europe,  the  type.  The  writer 
has  proposed  several  new  generic  groups  for  the  diverse  species 
usually  referred  to  Asterina,1  The  only  species  found  on  the  north- 
west coast  belongs  to  the  restricted  genus  Patina.  Other  genera 
are  found  farther  south,  and  may  occur  off  San  Diego. 

xThe  classification  adopted  (op.  cit.,  1913)  for  those  genera  most  nearly 
allied  to  Asterina  is  as  follows : 

A.  Disk  and  rays  depressed,  margins  thin,  the  plates  small. 
B$.  Principal  dorsal  plates  all  imbricated. 

C.  Ventral  or  interactinal  plates  bear  a  fan  of  two  to  eight,  usually  webbed 
spines. 

D.  Dorsal  plates  of  papular  areas  spinulated,  nearly  all  of  one  kind,  without 
large  groups  of  small  intervening  ossicles. 

E.  Two-bladed  pedicellariae  are  present  on  dorsal  plates.    Asterina  Nardo 
(restricted).    Type,  A.  gibbosa  (Penn.).    Europe. 

EE.  Pedicellarise  are  lacking.  Asterinides  Verrill.  Type,  A.  folium  (Lutk.). 
W.  Indies;  Bermuda;  Florida. 

DD.  Dorsal  plates  of  papular  areas  are  mostly  lobed  and  notched,  with  a 
curved  or  lunate  crest  bearing  fine  spinules;  between  them  are  clusters  of 
many  very  small  spinulated  ossicles. 

F.  Pedicellariae  are  lacking.    Patiria  Gray,  1840.    Type,  P.  coccinea  Gray. 
S.  Africa  (monotypic).    Not  Patiria  of  Perrier  nor  of  Sladen. 

FF.  Pedicellariae  are  present  on  the  dorsal  plates.  Papular  areas  differ- 
entiated. Enoplopatiria  Verrill.  Type,  E.  marginata  (Hupe)  =£.  brasiliensis 
(Lutk.).  Brazil;  W.  Indies;  W.  Africa. 

CC.  Ventral  plates  are  without  fans  of  spines. 

G.  Ventral  plates  have  a  single  spine;  sometimes  two.    Dorsal  plates  nearly 
as  in  Patiria.    Patiriella  Verrill.    Type,  P.  regularis  Ver.    New  Zealand. 

GG.  Ventral  plates  have  a  fascicle  or  cluster  of  spines. 

H.  Ventral  plates  have  a  fascicle  of  slender  spines.  Asterinopsis  Verrill. 
Type,  A.  penicillaris  (Lam.).  Indian  and  Pacific  Oceans;  Australia. 

HH.  Ventral  plates  have  a  central  cluster  of  many  short  spinules  or  gran- 
ules ;  rays  longer ;  rounded.  Callopatiria  Verrill.  Type,  C.  bellula  Sla. 

BB.  Principal  dorsal  plates  finely  spinulated,  not  all  imbricated;  thin,  trans- 
versely elongated,  with  many  small  intervening  ossicles.  Plates  rest  on  an 
alveolar  structure,  tubular  interradially  distally.  Ventral  plates,  toward  mouth, 
have  an  irregular  group  of  webbed  spines ;  distally  changing  to  a  regular  comb 
of  about  three.  Desmopatiria  Verrill.  Type,  D.  Aexilis  Ver.  Chile  (?). 

AA.  Disk  and  rays  not  much  depressed ;  rays  longer,  rounded ;  margins  not 
thin  and  subacute;  larger  dorsal  plates  not  all  imbricated.  Here  are  placed 
Paro^oiwa  Fisher.  Type,  P.  crassa  (Gray).  Allopatiria  Verrill.  Type,  A. 
ocellifera  (Gray). 


264  VERRILL 

Genus  Patiria  Gray. 
Type,  P.  coccinta  Gray,  1840. 

Patiria  GRAY,  1840;  Gray  (pars),  op.  cit.,  1866,  p.  16  (not  of  Perrier,  nor  of 

Sladen). 

Asterina  (pars)  PERKIER,  SLADEN,  FISHER,  and  others. 
Patiria  VERRILL,  op.  cit.,  1867;  Amer.  Journ.  Sci.,  vol.  xxxv,  pp.  480,  482, 

May,  1813. 

Disk  broad,  usually  pentagonal,  sometimes  hexagonal,  depressed, 
with  thin  margins  and  short  rays.  Dorsal  plates  unequal,  spinu- 
lated,  more  or  less  imbricated,  the  larger  ones  crescent-shaped,  with 
the  exposed  edges  rounded,  becoming  small  distally;  between  these 
are  many  much  smaller  rounded  ones.  Marginal  plates  small, 
scarcely  larger  than  the  adjacent  dorsals,  upper  ones  usually  some- 
what smaller;  both  with  marginal  spinelets.  Papulae  are  distributed 
over  most  of  the  upper  surface  of  the  rays  and  central  part  of  disk 
in  radial  rows.  Interactinal  plates  have  spines  in  small  combs,  often 
webbed.  The  larger  dorsal  ossicles  have  three  or  four  internal  lobes, 
by  which  they  are  connected  together.  The  distal  interradial  plates 
have  internal,  conical,  descending  processes  which  join  projections 
from  the  ventral  plates. 

Adambulacral  plates  have  a  furrow-comb  of  about  three  to  five 
webbed  spines,  and  a  group  of  two  to  five  spines,  in  a  row. 

Madreporic  plate  large,  dorsal,  much  nearer  to  the  center  than  the 
margin. 

PATIRIA  MINIATA  (Brandt)  Verrill.  ' 
Plate  vn,  figures   I,  2;  plate  cvni,  figures  I,  2   (varieties). 

Asterias  miniata  BRANDT,  Prodromus,  p.  68,  1835. 

Asteriscus  miniatus  STIMPSON,  op.  cit.,  vi,  p.  90,  1857. 

Patiria  miniata  VERRILL,  Trans.  Conn.  Acad.,  i,  pp.  234,  236,  1867  (distri- 
bution) ;  Amer.  Journ.  Sci.,  vol.  xxxv,  p.  482,  1913. 

Asterina  miniata  SLADEN,  op.  cit.,  p.  774,  1889.  Verrill,  American  Naturalist, 
XLIII,  p.  547,  fig.  2  (six-rayed),  September,  1909.  Fisher,  op.  cit,  19116, 
p.  254,  pi.  LVI,  figs.  6,  8;  pi.  LXI,  figs.  1-4;  pi.  LXII,  figs,  i,  2. 

Size  large ;  disk  rather  thick ;  rays  normally  five,  often  six,  trian- 
gular, rapidly  tapered,  about  as  broad  as  long.  Radii  of  a  large  five- 
rayed  specimen,  46  mm.  and  92  mm. ;  ratio,  1:2.  A  large  six- 
rayed  specimen  has  the  radii  44  mm.  and  85  mm. ;  ratio,  i :  1.93. 
The  disk  is  convex  or  swollen,  in  well  preserved  specimens,  along 
the  radial  areas,  but  usually  with  depressions  between,  in  large 
specimens. 

The  larger  dorsal  plates  along  the  median  radial  areas  of  the  disk 
and  ravs  are  prominent,  crescent-shaped,  covered  with  short,  rough 


SHALLOW-WATER   STARFISHES  265 

spinules.  Alternating  irregularly  with  these  are  many  small  oblong 
and  rounded  plates,  similarly  granulated.  Everywhere  in  the 
depressed  areas  between  the  larger  plates  there  are  many  lower  and 
much  smaller  ossicles,  unequal  in  size  and  form,  bearing  small 
clusters  of  small  granules.  The  larger  plates  form  pretty  regular 
radial  rows  medially  on  the  rays,  and  oblique  ones  on  their  sides. 

Toward  the  margin,  the  crescent-shaped  plates  become  smaller  and 
shorter,  and  near  the  margin  of  the  disk  all  become  rounded  or 
elliptical,  with  a  rosette  of  granules ;  while  on  the  sides  of  the  rays, 
distally,  they  become  elliptical  and  oblong,  closely  spinulated. 

The  marginal  edge  is  formed  by  the  two  regular  rows  of  mar- 
ginal plates,  which  are  only  a  little  larger  than  the  adjacent  dorsals, 
and  transversely  elliptical  in  form,  as  seen  covered  with  a  wreath  of 
spinules,  slightly  longer  than  those  of  the  dorsals.  The  lower  mar- 
ginals are  slightly  the  larger.  The  rows  are  separated  by  a  narrow 
but  well  defined  groove.  The  actinal  plates  form  large  triangular 
areas.  They  are  closely  arranged  in  many  alternate  rows.  Each 
bears  a  group  of  three  to  five  erect,  slightly  divergent,  partly  webbed 
spines,  which  become  much  longer  and  larger  adorally. 

The  adoral  spines  are  stout  and  obtuse,  about  three  or  four  pairs. 
The  epiorals  are  larger,  stout,  blunt,  connivent,  often  only  one  pair 
on  small  specimens,  and  then  they  resemble  spiniform  pedicellariae. 
On  larger  specimens  there  may  be  four  to  six  connivent  pairs,  and 
then  they  resemble  pectinate  pedicellariae. 

The  adambulacral  plates  bear  a  furrow-series  of  two  or  three 
slender,  parallel,  partly  webbed  spines  and  two  or  three  larger, 
longer,  erect,  blunt  spines  on  the  actinal  side. 

The  interactinal  plates  are  covered  with  a  thin  skin  that  forms 
delicate,  narrow,  ciliated  canals,  transverse  to  the  rays,  and  running 
out  from  between  .the  adambulacral  groups  of  spines. 

Color,  in  life,  commonly  orange,  varying  to  scarlet,  bright  red, 
dark  red  or  red-brown,  purple,  lemon-yellow,  etc. ;  sometimes  green- 
ish or  olive,  or  blotched  with  green. 

Its  range  extends  from  San  Diego  to  middle  Alaska.  I  have 
studied  specimens  from  many  localities.  It  is  abundant  on  the  coast 
of  California,  in  shallow  water  and  between  tides,  on  rocky  shores, 
at  Monterey  and  many  other  places.  Pacific  Grove  (W.  R.  Coe), 
many  large  examples;  San  Luis  Obispo  (Prof.  E.  T.  Nelson). 
Common  at  Vancouver  Island.  Numerous  large  specimens  were  col- 
lected at  Departure  Bay,  Vancouver  Island,  in  1909,  by  Prof.  John 
Macoun  and  party  of  the  Canadian  Geological  Survey.  Most  of 


266  VERRILL 

these  were  dark  purple,  but  some  were  orange.  Among  them  were 
two  six-rayed  specimens  and  one  very  large  one  with  seven  perfectly 
regular  rays.  Fisher  records  it  from  many  localities,  mostly  off  the 
California  coast,  from  low  tide  to  165  fathoms,  and  also  from  the 
Gulf  of  California,  north  of  La  Paz,  in  33  fathoms. 

VARIATIONS. 

Many  regular  six-rayed  examples  occur  of  full  size.  From  Pacific 
Grove  (coll.  Prof.  W.  R.  Coe),  there  is  also  a  four-rayed  specimen 
which  is  quite  regular,  except  that  one  ray  has  been  broken  off  and 
not  fully  restored.  A  regular  seven-rayed  specimen  has  been 
received  from  Vancouver  Island  (pi.  cvm,  fig.  i),  Canadian  Geologi- 
cal Survey.  Its  radii  measure  58  mm.  and  84  mm.  With  the  latter 
was  a  regular  six-rayed  specimen,  with  the  radii  48  mm.  and  72  mm. 

Suborder  VELATA  Perrier.    (See  p.  204.) 
Family  PTERASTERID&.  Perrier. 

Pterasterida  PERRIER,  op.  cit.,  1875. 

Pterasterida  (pars)  SLADEN,  Voy.  Challenger,  vol.  xxx,  p.  xxxvii,  p.  468,  1889. 
Fisher,  op.  cit.,  191  ib,  p.  343. 

This  is  one  of  the  most  peculiar  groups  of  starfishes  hitherto  dis- 
covered. It  shows,  in  general  characters,  a  remarkably  high  degree 
of  specialization  not  found  in  any  other  group.  Most  of  the  genera 
and  species  are  from  the  deep  sea. 

Disk  usually  plump ;  rays  five  to  eight,  rarely  nine ;  upper  surface 
covered  by  a  supradorsal  membrane,  supported  by  the  tips  of  long, 
slender,  divergent,  often  webbed,  paxillary  spinules  and  pierced  by 
small  concentric  pores  or  "  spiracles,"  often  closed  and  invisible  in 
preserved  specimens,  and  usually  with  a  larger  central  osculum. 
Beneath  this  membrane  is  a  "nidamental"  cavity  or  gonocodium, 
traversed  by  the  columnar  pseudopaxillae,  and  containing  the 
papulae,  which  are  often  branched.  In  this  cavity  the  eggs  are 
retained,  and  also  the  young,  till  they  assume  the  adult  form  and 
considerable  size. 

The  dorsal  skeletal  ossicles  are  lobed  or  cruciform,  loosely  reticu- 
lated. The  adambulacral  spines  usually  form  transverse  webbed 
combs  or  fans;  not  webbed  in  Hymenasterinse.  Series  of  slender, 
divergent  spines,  more  or  less  appressed,  and  attached  to  the  under 
surface,  or  imbedded  in  it,  and  usually  webbed  to  the  adambulacral 
fans  in  Pterasterinse,  are  always  present.  These  peculiar  spines, 
called  "  actino-lateral  spines "  by  Sladen,  should  rather  be  called 


SHALLOW-WATER   STARFISHES  267 

retroambulacrals,  or  outer  adambulacrals,  for  they  are  attached  to 
the  outer  end  of  the  adambulacrals.  They  are  sometimes  short  and 
do  not  reach  the  margin,  except  distally,  on  the  rays;  but  in 
Pteraster  and  some  other  genera  their  tips  usually  reach  the  margin 
of  the  disk.  Between  the  bases  of  these  there  is  usually  a  small 
slit  or  pore  (actinal  spiracle  or  "  segmental  aperture"),  furnished 
with  a  calcareous  valve. 

Interactinal  plates  are  lacking.  Pedicellariae  have  not  been  found. 
Jaws  have  a  series  of  adoral  spines,  usually  webbed.  Epioral  spines 
of  large  size,  in  one  to  three  pairs,  often  without  webs,  stand  per- 
pendicularly to  the  jaw ;  those  of  one  pair  are  sometimes  specialized 
and  partially  hyaline.  Ambulacral  feet  are  large,  in  two  to  four  rows, 
with  large  apical  suckers. 

The  supradorsal  membrane  may  be  thin  and  translucent,  with 
thin  muscular  fibers,  or  thick  and  muscular;  it  may  contain  calca- 
reous spicules,  muscular  and  cartilaginous  fibers  in  the  form  of  a  net- 
work, and  often  abundant  mucous  glands.  The  copious  mucus  is 
phosphorescent  in  some  cases  (Diplopteraster) . 

The  intestine  and  anal  pore  are  well  developed.  Sexes  are  alike 
externally. 

Subfamily  HYMEN ASTERINJE  Verrill,  nov. 

The  genera  Hymenaster  and  Cryptaster,  and  their  allies,  should 
form  a  distinct  subfamily  Hymenasterinae,  differing  from  Pteraster- 
inse  in  lacking  webbed  combs  of  adambulacral  spines  and  in  other 
characters.  It  is  a  characteristic  deep-sea  group. 

Pythonaster  Sladen  does  not  belong  to  this  family.  It  has  no 
supradorsal  membrane,  actinomarginal  spines,  nor  actinal  spiracles. 

Subfamily  PTERASTERIN1E. 
Genus  Pteraster  Mtiller  and  Troschel. 

Pteraster  MULLER  and  TROSCHEL,   Syst.  Aster.,  p.   128,   1842.     Sladen,  op. 

cit.,  p.  470,  1889.     Fisher,  op.  cit.,  191  ib,  p.  344;  analytical  table  of  all 

known  species,  pp.  368-370. 
Hexaster  PERKIER,  Comptes  rendus,  cxn,  p.   1227,  1891   (type,  H.  obscurus 

Perrier). 
Temnaster  VERRILL,  Proc.  U.  S.  Nat.  Mus.,  xvn,  p.  275,  1894  (type*  T.  hexactis 

Verrill  =  H.  obscurus  Perrier). 

Disk  large ;  rays  five  to  eight ;  ambulacral  grooves  turn  up  distally, 
more  or  less,  reaching  to  the  upper  side  of  the  rays. 

Adambulacral  spines  form  equal,  webbed  combs;  retroambulacral 
or  actinomarginal  spines  closely  appressed,  ending  in  a  fringe  or 


268  VERRILL 

fold.  One  pair  of  free  epioral  spines;  these  are  often  translucent 
distally.  Supradorsal  membrane  furnished  with  a  central  contractile 
osculum,  guarded  by  five  groups  of  webbed,  projecting  spines.  It 
usually  contains  irregular  muscular  fibers  for  its  support;  muscular 
bands  usually  do  not  form  a  regular  network  and  are  generally  very 
inconspicuous  externally.  Usually  two  rows  of  ambulacral  feet,  or 
nearly  four-rowed,  by  crowding,  in  P.  marsippus. 

PTERASTER  TESSELATUS  Ives. 

Plate  xxxn,  figures  i,  2;  plate  LXXXVI,  figures  4-4^  (details);  plate  xcvn, 

figure  i. 

Pier  osier  tesselatus  IVES,   Proc.  Acad.   Nat.   Sci.   Philadelphia,   for   1888,  p. 

421 ;  and  1889,  p.  169.    Fisher,  op.  cit.,  19116,  p.  359,  pi.  civ,  figs.  1-5. 
Pier  osier  reticulatus  VERRILL,  op.  cit.,  19096,  p.  555   (an  accidental  error  in 

spelling) . 

A  large,  plump,  five-rayed  species,  with  wide,  tapered,  subacute 
rays  of  short  or  moderate  length,  and  with  high,  rounded  margins  to 
the  disk. 

The  ambulacral  grooves  are  turned  up  a  little  at  the  end  of  the 
rays,  so  that  the  distal  and  apical  plates  are  dorsal.  Submarginal 
fold  narrow. 

Radii  of  the  largest  (dry)  specimen  from  Puget  Sound  (Prof. 
Trevor  Kincaid)  are  42  mm.  and  70  mm.  to  75  mm. ;  height,  50  mm. 
Radii  of  one  from  off  Sidney,  British  Columbia,  are  28  mm.  and 
40  mm. ;  height,  35  mm.  The  alcoholic  specimen  figured,  from  the 
Harriman  Expedition,  is  larger,  the  greater  radius  measuring  90  mm. 
or  more. 

The  dorsal  surface  is  covered  with  a  strong,  thick,  supradorsal 
membrane,  through  which  the  spinules  of  the  pseudopaxillae  usually 
show  only  as  very  small  points,  arranged  in  a  complex  reticulated 
pattern.  But  in  the  largest  dry  specimens  they  appear  as  small,  more 
elevated,  obtuse  spines  or  tubercles,  sometimes  showing  a  stellate 
arrangement,  but  more  often  closely  reticulated  or  areolate.  The 
dorsal  osculum  is  relatively  rather  small,  with  elevated  spinose  mar- 
gins, the  spines  not  very  long,  rather  stout,  the  points  not  pro- 
jecting beyond  the  webs. 

The  adambulacral  fans  of  spines  consist  of  about  three  or  four 
outer,  rather  long,  nearly  equal,  slightly  tapered  spines,  and  two 
much  smaller  inner  ones,  of  which  the  innermost  is  only  about  one- 
fourth  as  long  as  the  outer  ones,  and  the  next  one  is  about  half  their 
length.  In  some  large  specimens  these  spines  become  clavate. 


SHALLOW-WATER   STARFISHES  269 

The  appressed  retroambulacral  or  "  actino-marginal "  spines  are 
unusually  short  and  stout,  a  little  flattened  and  enlarged  distally, 
and  almost  twice  as  long  as  the  adambulacrals ;  but  they  do  not  reach 
a  third  of  the  distance  to  the  margin  in  the  middle  of  the  interradial 
areas,  so  that  the  dorsal  spinulation  extends  well  onto  the  under  side, 
making  the  margins  thick  and  swollen.  They  end  at  a  strong,  free 
fold  of  the  membrane,  which  only  becomes  marginal  near  the  ends 
of  the  rays.  The  peractinal  spiracles  or  pores  are  narrow  slits,  with 
narrow,  elongated,  somewhat  crescent-shaped  valves.  Adoral  spines 
form  about  five  graded  pairs,  completely  webbed ;  the  terminal  ones 
are  rather  stout;  the  distal,  very  slender.  Epiorals  large,  tapered, 
translucent  distally.  The  dorsal  spiracles  are  small  and  numerous, 
but  in  dry  specimens  and  some  alcoholics  they  are  invisible,  owing 
to  strong  contraction. 

This  species  is  known  from  Bering  Sea  to  Puget  Sound.  My 
specimens  are  from  Yakutat  (Professor  Ritter)  and  Sitka  (W.  R. 
Coe),  Harriman  Expedition;  Puget  Sound  (Professor  Kincaid)  ; 
off  Sidney  Islands,  May  10,  1896  (Prov.  Mus.  B.  C.)  ;  British  Colum- 
bia (Canada  Geological  Survey).  Mr.  Ives  recorded  it  from  Marmot 
Island,  Alaska  (type  locality).  Fisher  recorded  it  from  Bering  Sea, 
off  Pribilof  Islands ;  off  Shumagin  Islands ;  off  Unalaska,  and  various 
other  Alaskan  localities,  from  low  water  to  150  fathoms;  off  Van- 
couver Island,  in  238  fathoms;  Straits  of  Fuca,  114  fathoms;  near 
Port  Townsend,  15  to  39  fathoms;  etc.  Some  of  his  specimens  were 
much  larger  than  the  one  described  above.  He  gives  the  radii  of  one 
specimen  as  54  mm.  and  88  mm. ;  ratio,  1 :  1.63. 


PTERASTER  TESSELATUS  ARCUATUS  Fisher. 
Pteraster  tcsselatus  arcuatus  FISHER,  op.  cit,  191  ib,  p.  363,  pi.  cm,  fig.  I. 

Similar  to  P.  tesselatus,  but  arcuately  pentagonal  in  form,  with 
thick  supradorsal  membrane,  without  signs  of  reticulation ;  spiracula 
sunken  in  irregular  creases.  Radii,  32  mm.  and  46  mm.;  ratio, 
i :  1.44. 

Color  in  life,  above,  bluish  gray  or  dull  heliotrope  purple,  mottled 
with  fawn-color,  with  dark  interradial  and  lighter  radial  areas,  and 
dark  around  margins ;  interradial  areas  below,  dull  heliotrope  purple ; 
marginal  membrane,  vinaceous  buff  (teste  Fisher). 

Monterey  Bay,  in  46  to  56  fathoms  (Fisher). 

This  appears  to  be  the  same  as  the  next,  grown  to  larger  size. 


27O  VERRILL 

PTERASTER  TESSELATUS  HEBES  Verrill. 

Plate  xcvi,  figures  i,  2  (type,  enlarged). 
Pier  osier  hebes  VERRILL,  Amer.  Journ.  Sci.,  xxvni,  p.  61,  1909. 

Disk  plump  and  relatively  large,  the  five  rays  being  very  short  and 
blunt,  with  the  ambulacral  grooves  and  plates  turned  upward  and 
reflexed  upon  the  upper  surface  nearly  to  the  base  of  the  rays,  or 
about  even  with  the  shallow  interradial  angles.  Radii,  22  mm.  and 
28  mm. ;  ratio,  1 :  1.28. 

The  central  dorsal  oscule  is  well  developed,  surrounded  with 
slender,  webbed,  projecting  spines,  in  five  groups  of  eight  to  ten  each. 
The  dorsal  surface  is  covered  with  a  multitude  of  crowded  slender 
spinules,  which  project  above  the  marsupial  membrane  and  give 
almost  the  appearance  of  velvet  pile;  but  in  some  places  they  form 
more  or  less  evident  divergent  stellate  clusters  of  twelve  to  twenty 
spinules.  Seen  from  within,  these  spinules  are  slender,  2  mm.  to 
3  mm.  long,  very  divergent,  supported  by  slender  columnar  paxillse. 

The  ambulacral  grooves  are  broad  and  shallow.  The  ambulacral 
plates  are  somewhat  bilobed  at  the  inner  ends,  and  distally  are  some- 
what imbricated.  The  adambulacral  spines  are  long  and  slender, 
about  five  or  six  in  a  transverse  row,  of  which  the  two  inner  ones  are 
very  small  and  slender,  not  half  as  long  as  the  outer  ones,  of  which 
there  are  three  or  four,  about  3.5  mm.  long. 

The  appressed  actino-marginal  spines  are  distinctly  longer,  about 
twice  as  stout,  and  blunt  proximally  on  the  ray,  but  distally,  on  the 
upturned  part,  where  they  are  crowded,  they  become  about  equal  in 
length  to  the  adambulacrals  and  scarcely  larger;  those  near  the 
interradial  angles  are  flattened  and  enlarged  distally;  the  valves  at 
the  peractinal  pores  between  their  bases  are  very  acute,  small,  and 
slender,  as  seen  edgewise,  but  when  removed  they  are  acute-triangu- 
lar, and  curved  proximally. 

Departure  Bay,  British  Columbia,  23  fathoms,  mud  and  sand,  1908 
(C.  H.  Young,  Canadian  Geological  Survey).  One,  dry  and  stuffed. 

The  jaws  and  entire  oral  region  have  been  destroyed  by  the  pre- 
parator.  Therefore  its  relations  to  some  of  the  other  species  are 
uncertain.  It  seems  to  be  nearest  to  P.  tesselatus,  of  which  it  is  prob- 
ably only  a  variety ;  some  of  the  differences  are  probably  due  to  imma- 
turity. The  mode  of  preparation  and  drying  may  have  increased 
the  size  of  the  disk  considerablv. 


SHALLOW- WATER   STARFISHES  271 

PTERASTER  MULTISPINUS  Clark. 

Ptet -aster  multispinus  H.  L.  CLARK,  Proc.  Boston  Soc.  Nat.  Hist.,  xxix,  No.  15, 
p.  326,  pi.  in,  figs.  I,  2,  1901.  Fisher,  op.  cit,  19116,  p.  359  (description 
copied  from  Clark). 

The  type  specimen,  which  I  have  studied  in  the  collections  of 
Columbia  University,  is  a  small  five-rayed  specimen,  with  short, 
tapered,  subacute  rays,  broader  than  long,  with  the  small  tips  turned 
up  a  little  at  the  ends.  Radii,  18  mm.  and  27  mm.;  ratio,  i:  1.5. 
It  is  probably  young;  it  is  preserved  in  alcohol. 

Disk  a  little  convex,  covered  with  small,  numerous,  slightly  pro- 
jecting spinules;  there  is  no  defined  margin,  the  borders  being 
rounded  and  the  dorsal  membrane  continuous  to  the  under  side  in  the 
interradial  areas.  The  dorsal  membrane  is  rather  firm,  with  very 
small  spiracular  pores,  inconspicuous  except  distally  on  the  rays. 
Pseudopaxillae  are  hour-glass  shaped,  rather  high,  bearing  eight  to 
ten,  or  often  more,  spinules.  Oscular  opening  rather  large,  sur- 
rounded by  five  clusters  of  webbed  spines,  about  six  in  each  group, 
the  central  ones  larger. 

Ambulacral  feet  in  two  rows.  Adambulacral  fans  of  spines  are 
oblique,  each  with  five  or  six  spines,  webbed  to  the  tips.  The  inner 
one  is  much  the  shortest;  the  outer  three  are  longest.  Appressed 
retroambulacral  spines  are  mostly  short.  Therefore  there  is  no 
marginal  free  edge  with  a  fringe  of  spine-tips  along  the  rays,  such 
as  is  conspicuous  in  P.  militaris  and  many  other  species,  but  some 
of  the  spines  in  the  interradial  areas  are  rather  long  and  slender. 

Peroral  spines  slender,  six  on  each  angle  of  the  jaw,  webbed 
together.  Epiorals  two,  rather  large,  slightly  swollen  distally  at 
the  translucent  part;  tips  acute.  Peractinal  or  segmental  pores 
with  a  narrow  spiniform  valve,  as  seen  edgewise,  in  natural  position. 

(  ?)  Puget  Sound  (Columbia  University  Expedition,  1897). 

This  looks  very  much  like  a  young  specimen  of  P.  tesselatus, 
younger  than  any  of  the  latter  that  I  have  seen.  The  apparent  lack 
of  numerous  spiracles  on  the  disk  is  due  to  the  strong  contraction  of 
the  integument  in  alcohol.  It  often  occurs  in  P.  tesselatus  also. 

PTERASTER  GRACILIS  (Clark)  Verrill. 

Retaster  gracilis  H.  L.  CLARK,  Proc.  Boston  Soc.  Nat.  Hist.,  xxix,  p.  326, 

pi.  in,  figs.  3,  4,  1897. 
Pter aster  gracilis  VERRILL,  op.  cit.,  1909,  p.  555.    Fisher,  op.  cit.,  1911,  p.  349 

(description  copied  from  Clark). 

This  is  a  five-rayed  species,  of  which  I  have  studied  the  type  at 
Columbia  University.  It  has  the  radii  about  9  mm.  and  18  mm. 


272  VERRILL 

The  rays  are  about  as  long  as  broad,  acute,  slightly  turned  up  at 
tips.  It  is  evidently  young.  It  was  preserved  in  alcohol. 

Dorsal  surface  reticulated,  due  mainly  to  the  slender,  divergent 
spinules  of  the  pseudopaxillae.  These  spinules,  which  form  stellate 
groups  of  six  to  twelve  or  more  on  the  slender  paxillae,  are  long 
and  thin.  Oscular  opening  small,  surrounded  by  five  groups  of 
small  spines. 

Adambulacral  combs  of  spines  contain  four  to  six,  most  often 
five.  Of  these  the  inner  one  is  very  small ;  then  three  or  four  much 
longer,  subequal,  central  ones.  Retroambulacral  spines  longer  and 
larger,  horizontally  webbed,  and  partly  erect.  The  horizontal  web 
has  a  free  edge  parallel  with  the  ambulacral  groove.  Peractinal 
spiracles  small,  with  a  narrow  valve,  free  laterally. 

Jaws  with  five  webbed  and  graded  spines,  the  apical  ones  much 
the  larger ;  lateral  ones  small  and  slender.  Epioral  spines  two,  large ; 
distal  half  nearly  transparent;  base  swollen,  white,  opaque;  tips 
tapered,  slender,  acute. 

Ambulacral  feet  in  two  rows. 

(?)  Puget  Sound  (Columbia  University  Expedition). 

This  may  prove  to  be  the  young  of  P.  tesselatus,  when  specimens 
of  intermediate  sizes  can  be  studied.  The  smallest  typical  example  of 
the  latter  known  to  me  is  80  mm.  in  diameter.  They  agree  in  the 
shortness  of  the  retroambulacral  spines,  which  do  not  reach  the 
margin  of  the  disk  in  either  case,  but  end  in  a  raised  submarginal 
web,  parallel  with  the  groove.  The  differences  that  exist  may  very 
likely  be  due  to  age  and  mode  of  preservation.  I  see  no  reason  for 
referring  this  species  to  Retaster.  The  reticulating  lines,  mentioned 
by  Clark,  seem  to  me  to  be  merely  very  slender,  divergent  paxillary 
spinules,  not  muscular,  nor  cartilaginous  fibers.  Such  muscular  fibers 
as  were  noticed  by  me  were  very  feeble  and  irregular. 

PTERASTER  MILITARIS  (Miiller)  Miiller  and  Troschel. 

Asterias  militaris  MULLER,  Zool.  Dan.  Prod.,  p.  234,  1776.    Rathke,  Zool.  Dan., 

iv,  p.  14,  pi.  ex  xxi,  1806. 
Pteraster  militaris  MULLER  and  TROSCHEL,  Syst.  Aster.,  Suppl.,  p.  128,  pi.  vi, 

fig.  I,  1842.    Stimpson,  Invert.  Grand  Manan,  p.  15,  1853.    M.  Sars,  Overs. 

Norges  Echinod.,  p.  48,  pi.  in,  figs.  8,  9,  pi.  iv,  figs.  4-6,  1861.     Duncan 

and  Sladen,  op.  cit,  p.  46,  pi.  in,  figs.  13-16.     Danielssen  and  Koren,  op. 

cit,  p,  70,  pi.  xin,  figs.  18,  19,  1884.    Verrill,  Expl.  by  the  Albatross,  p. 

541,  pi.  xin,  fig.  35,  1885 ;  Distrib.  Echinod.,  p.  202,  1895. 

Pteraster  militaris  FISHER,  op.  cit,  1910,  p.   167;  op.  cit,  191  ib,  p.  346,  pi. 
xcvin,  figs.  I,  2. 

This  well-known  Arctic  and  North  Atlantic  species  has  been 
described  by  Mr.  Fisher  from  several  stations  off  the  Pacific  coast, 


SHALLOW-WATER   STARFISHES  273 

all  from  northern  Alaska  and  Bering  Sea,  in  41  to  344  fathoms, 
except  one,  from  the  Straits  of  Fuca,  in  100  fathoms. 

It  is  found  on  both  coasts  of  the  North  Atlantic,  in  10  to  530 
fathoms.  On  the  East  American  side  it  ranges  south  to  Cape  Cod. 
Common  in  the  Bay  of  Fundy,  in  10  to  50  fathoms.  On  the 
European  side  it  extends  south  to  Norway,  Scotland,  and  England. 
Common  in  Barents  Sea,  Spitzbergen,  Kara  Sea,  Greenland,  etc.  It 
is  evidently  circumpolar  in  distribution. 

Some  of  Dr.  Fisher's  specimens  were  unusually  large.  He  gives 
the  radii  of  one  as  33  mm.  and  75  mm. ;  ratio,  1 : 2.2. 

PTERASTER  PULVILLUS  M.  Sars. 

Pteraster  puhillus  SARS,  Overs.  Norges  Echinod.,  p.  62,  pi.  vi,  figs.  14-16,  pis. 
vii,  vni,  1861.  Verrill,  Amer.  Journ.  Sci.,  xvi,  p.  371,  1878;  Distrib. 
Echinod.,  p.  202,  op.  cit.,  1895.  Fisher,  op.  cit.,  191  ib,  p.  358,  pi.  en, 
figs.  2,  4. 

Two  specimens  of  this  Arctic  and  North  Atlantic  species  are 
described  by  Fisher  from  off  Bering  Island,  in  Bering  Sea,  in  72 
fathoms. 

On  the  eastern  American  coast  it  is  rare.  It  has  been  taken  off  the 
coasts  of  New  Hampshire  and  Maine,  and  in  the  Bay  of  Fundy,  in 
20  to  in  fathoms,  and  northward  to  the  Grand  Banks.  It  has  been 
recorded  from  the  Arctic  Ocean,  Barents  Sea,  Kara  Sea,  Spitz- 
bergen, etc.  On  the  European  coast  it  ranges  southward  to  Norway. 

PTERASTER   MARSIPPUS   Fisher. 

Pteraster  marsippus  FISHER,  op.  cit.,  1910,  p.  168;  op.  cit.,  191  ib,  p.  352,  pi. 
c,  fig.  I ;  pi.  ci,  fig.  2 ;  pi.  cxv,  fig.  4 ;  pi.  cxvi,  fig.  I. 

A  large,  stellate,  five-rayed  species.  Radii,  40  mm.  and  100  mm. ; 
ratio,  i :  2.5. 

Dorsal  membrane  thick,  smooth,  without  many  prominent  paxillary 
protrusions,  but  with  a  few  calcareous  spicular  deposits.  It  has  so 
few  or  small  spiracles  that  they  were  not  visible,  as  preserved. 

Adambulacral  combs,  proximally,  have  five  spines  with  strong  web 
and  sacculus  extending  beyond  tips.  A  longitudinal,  narrow  web 
between  successive  combs,  at  the  outer  border. 

Tube-feet  crowded  so  as  to  appear  to  be  in  four  rows,  except  in 
young. 

Bering  Sea,  in  52  to  351  fathoms,  southward  to  the  Aleutian 
Islands. 

This  species,  in  the  partially  four-ranked  arrangement  of  the  tube- 
feet,  approaches  Diplopter aster. 
19 


274  VERRILL 

PTERASTER  OBSCURUS  (Perrier)  Doderlein. 

Hexaster  obscurus  PERKIEK,  Mem.  Soc.  Zool.  France,  iv,  p.  267,   1891 ;  Res. 

Camp.  Sci.,  xi,  p.  41,  pi.  in,  figs,  i,  la,  1896. 

Pterastcr  (Temnaster)  hexactis  VERRILL,  Proc.  Nat.  Mus.,  xvn,  p.  275,  1894. 
Temnaster  hexactis  VERRILL,  Amer.  Journ.  Sci.,  XLJX,  p.  202,  1895. 
Hexaster  obscurus  VERRILL,  Revision,  Trans.    Conn.  Acad.,  x,  p.  271,  1899. 
Pteraster  octaster  VERRILL,   Amer.   Journ.   Sci.,   xxvm,   p.   61,   fig.    i,    1909 

(variety). 
Pteraster  obscurus  FISHER,  op.  cit.,  1911^,  p.  363,  pi.  cv,  figs.   1-4;  pi.  cvi, 

figs,  i,  2. 

The  original  description  of  P.  hexactis  was  as  follows : 

"  Disk  broad,  very  high,  evenly  convex,  with  a  rather  large  central 
opening  surrounded  by  circles  of  prominent,  imbricated,  and  webbed 
spines.  Rays  six,  short,  broad,  tapered  to  blunt  tips,  their  lateral 
margins  convex.  Lesser  to  greater  radii,  about  as  i  to  1.5.  Lesser 
radii,  22  mm. ;  greater  radii,  32  mm.  to  35  mm.,  in  the  alcoholic 
specimen ;  height  of  disk,  30  mm. 

"  The  surface  of  the  disk  is  covered  with  very  numerous  small 
spinules,  covered  more  or  less  completely  with  a  thick  skin-like 
membrane,  and  arranged  in  irregular,  divergent  groups. 

"  The  integument  between  the  spinules  is  thick,  smooth,  firm,  and 
everywhere  perforated  by  numerous  very  small,  round  pores. 

"  In  each  interradial  region  there  is  a  narrow,  radiating  groove, 
lined  with  thick  naked  integument,  destitute  both  of  spinules  and 
pores,  but  showing  a  wrinkled  surface.  These  grooves  commence  at 
about  one-fourth  the  distance  from  the  dorsal  center  to  the  margin. 
In  some  cases  there  is  only  a  small  slit-like  opening  in  the  upper  end 
of  the  groove,  communicating  with  the  space  beneath  the  dorsal 
membrane,  but  in  some  of  the  interradii  the  slit  is  much  larger  and 
longer,  reaching  nearly  or  quite  to  the  margin,  and  communicates 
with  a  large  marsupial  pouch,  containing  well-formed  young,  some 
of  which  were  in  the  act  of  escaping  when  preserved.  Apparently 
the  slit-like  openings  are  formed,  or  at  least  much  enlarged,  when  the 
young  are  ready  to  come  forth,  and  after  their  birth  the  edges  of  the 
slits  may  become  again  united. 

"  The  dorsal  spines  or  pseudopaxillae  beneath  the  integument  are 
large,  stout,  rather  long,  and  surmounted  with  a  large  divergent 
group  of  long,  slender  spinules.  In  the  interradial  region,  within 
the  marsupial  pouch,  there  is  a  group  of  several  lobed  or  branched 
papulae  at  the  base  of  each  paxilliform  spine.  The  large  spines 
situated  along  each  side,  within  these  cavities,  have  rudimentary 
spinules  at  the  summit,  which  do  not  reach  the  outer  membrane,  so 


SHALLOW-WATER   STARFISHES  275 

that  they  stand  free  within  the  cavity,  thus  leaving  the  membrane 
unsupported  along  the  slits.  On  the  ventral  side  the  rays  are  nearly 
flat,  and  the  disk  around  the  mouth  is  deeply  concave. 

"  Each  ray  is  broadest  at  the  margin  of  the  disk.  The  transverse 
adambulacral  combs  are  numerous  and  covered  with  a  thick,  firm 
skin,  which  entirely  conceals  the  spines  in  alcoholic  specimens.  On 
the  broadest  part  of  the  ray,  opposite  the  margin  of  the  disk,  there 
are  mostly  four,  rarely  five,  spines  of  moderate  length  in  each  comb ; 
of  these  the  one  next  the  groove  is  somewhat  shorter  than  the  two  or 
three  which  succeed  it,  while  the  outermost  is  still  shorter  and 
directed  more  outward,  so  that  the  group  has  a  somewhat  rounded, 
but  not  very  elevated,  scolloped  margin,  the  membrane  receding 
somewhat  between  the  points  of  the  spines.  The  spines,  when 
exposed,  are  rather  slender,  flattened,  rough,  and  truncate  at  the  flat 
tip;  beyond  the  outer  spine  the  web  rapidly  becomes  less  elevated 
and  each  comb  lies  somewhat  obliquely  over  the  one  next  beyond  it, 
and  becomes  only  a  slightly  elevated  broad  fold  before  reaching  the 
margin.  These  folds  entirely  conceal  the  appressed,  outer  adambu- 
lacral spines,  which  extend  to  the  margins  of  the  rays,  but  project 
very  little  if  at  all  beyond  it,  so  that  the  margin  is  merely  crenulated, 
or  divided  into  small  blunt  lobes  by  slight  notches.  The  spiracles, 
between  the  outer  ends  of  the  webbed  adambulacral  spines,  are  ovate 
pores,  sometimes  closed  by  an  ovate  operculum;  but  in  other  cases 
they  are  occupied  by  a  small  group  of  two  or  three  short,  papula-like 
organs. 

"  The  jaws  are  surrounded  by  a  marginal  group  of  long,  slender, 
webbed  spines,  of  which  there  are  about  four  or  five  on  each  side; 
the  two  innermost  are  somewhat  the  larger.  On  the  actinal  side  of 
the  jaws  there  are  also  two  much  larger,  isolated  epioral  spines,  one 
on  each  plate ;  these  are  entirely  covered  by  a  thick  skin ;  when  this 
is  removed  the  spine  is  flattened,  tapered,  and  blunt  at  the  tip,  with 
a  rough  surface,  but  not  hyaline. 

"  Color,  in  alcohol,  dull  purple. 

"  The  ambulacral  feet  are  large  and  in  two  regular  rows." 

The  specimen  described  above  as  the  type  of  P.  hexactis  was 
taken  off  Newfoundland,  in  57  fathoms,  N.  lat.  43°  05'.  The  type 
of  obscurus  Perrier  was  from  the  Banks  of  Newfoundland,  in  155 
meters.  The  few  Atlantic  specimens  have  all  been  six-rayed,  except 
one  seven-rayed  specimen.  It  has  also  been  taken  at  Spitzbergen  and 
Greenland. 


276  VERRILL 

Professor  Fisher  records  it  from  a  number  of  localities  in  Bering 
Sea,  from  Bering  Strait  to  Bering  Island  and  Unalaska,  and  off 
Kamchatka,  in  17  to  85  fathoms.  I  have  also  had  six-rayed  speci- 
mens from  the  same  region. 

He  has  found  that  in  Bering  Sea  it  occurs  with  six  to  nine  rays, 
but  most  frequently  with  six  rays.  The  only  nine-rayed  specimen 
was  a  young  one  taken  from  the  marsupial  pouch  of  an  eight-rayed 
specimen.  One  of  my  eight-rayed  specimens  also  contained  a  nine- 
rayed  young  one.  Some  of  his  specimens  were  much  larger  than 
the  one  described  above.  The  largest  six-rayed  specimen  had  the 
radii  43  mm.  and  71  mm. ;  ratio,  1 : 6.5.  The  largest  seven-rayed 
one  had  the  radii  56  mm.  and  82  mm. ;  ratio,  1 : 6.  His  large 
specimens  had  five  or  six  spines  in  the  adambulacral  combs,  proxi- 
mally.  He  did  not  find  genital,  interradial  slits,  showing  that  they 
are  temporary,  for  the  escape  of  the  young,  as  suggested  above. 


PTERASTER  OBSCURUS,  var.  OCTASTER  Verrill. 

Pteraster  octaster  VERRILL,  Amer.  Journ.   Sci.,  xxvm,  p.  61,  fig.   i,   1909. 
Fisher,  191  ib,  pi.  cvi,  fig.  I. 

The  Pacific  form  with  eight  or  nine  rays,  most  commonly  eight, 
seems  worthy  to  retain  the  name  octaster,  as  a  variety.  The  original 
description  was  as  follows : 

"  Disk  large  and  plump ;  margins  well  defined  by  points  of  the 
actino-marginal  spines ;  rays  eight,  short,  about  as  wide  as  long,  sub- 
acute  ;  the  ambulacral  grooves  turn  up  but  little  at  the  tips.  Radii  of 
the  largest  example,  20  mm.  and  30  mm. 

"  Dorsal  surface  covered  with  a  thick  membrane  through  which 
the  tips  of  the  spinules  show  but  little  as  pretty  uniformly  scattered 
points;  in  alcohol  they  form  the  apex  of  small,  low,  conical,  fleshy 
elevations.  Central  oscule  small,  in  alcohol  inconspicuous,  its  short 
spines  covered  by  a  soft  membrane.  Ambulacral  feet  large,  in  two 
rows. 

"  Adambulacral  spines  form  combs  of  five  or  six  spines,  of  which 
the  innermost  is  much  smaller  and  more  slender  than  the  rest,  which 
are  rather  stout,  tapered,  subacute,  divergent ;  the  outer  ones  longer ; 
the  outermost  appressed  to  the  surface.  Epioral  pair  of  spines  long 
and  rather  stout,  tapered,  translucent  distally.  The  interradial  areas 
are  narrow,  with  rows  of  long,  stout,  imbedded  actino-marginal 
spines,  the  ends  of  which  project  a  little  at  the  margin  of  the  disk. 


SHALLOW-WATER   STARFISHES  277 

"  Four  specimens  were  sent  to  me  by  the  U.  S.  National  Museum. 
Three  were  from  Bering  Island,  collected  by  Dr.  Stejneger  and  Mr. 
N.  Grebnitsky,  in  1888.  One  was  from  Kamchatka,  collected  by 
N.  Grebnitsky." 

Dr.  Fisher  records  the  eight-rayed  form  from  Bering  Strait,  17 
fathoms;  Bering  Island;  and  from  the  Pribilof  Islands,  in  26 
fathoms. 

He  states  that  he  has  taken  seven-rayed,  eight-rayed,  and  nine- 
rayed  young  from  the  gonocodium  of  a  single  eight-rayed  specimen ; 
and  six-rayed  young  from  a  seven-rayed  specimen.  This  shows  that 
the  number  of  rays  is  of  no  great  importance  in  this  species.  The 
young,  when  they  leave  the  pouch,  are  from  12  mm.  to  20  mm.  in 
diameter  (teste  Fisher). 

One  of  the  eight-rayed  specimens  described  by  me  was  in  the 
act  of  giving  birth  to  a  nine-rayed  young  one,  about  12  mm.  in 
diameter.  In  that  specimen  the  interradial  slits  were  well  marked, 
with  smooth  edges  (not  ruptured),  not  only  where  there  were  young 
beneath,  but  also  in  the  interradial  areas  where  there  were  no  young. 
The  arrangement  of  the  paxillary  spines  in  the  interradial  areas,  is 
such  as  to  facilitate  the  formation  of  the  slits.  Whether  there  is 
a  sexual  difference  in  this  respect,  I  do  not  know. 

The  species  of  this  genus  might  be  thought  hermaphrodite,  for  it 
is  rather  unusual  to  find  adult  specimens  that  do  not  carry  young. 
But  Professor  Fisher  has  dissected  a  large  specimen  of  P.  jordani 
which  proved  to  be  a  male,  not  differing  externally  from  the  females. 
Probably  the  males  are  fewer  in  number  than  the  females. 

Genus  Pterasterides  Verrill. 
Type,  Pteraster  aporus  Ludwig. 
Pterasterides  VERRILL,  Amer.  Naturalist,  XLIII,  p.  547,  September,  1909. 

Differs  from  Pteraster  in  being  destitute  of  a  notable  central 
dorsal  oscule  and  in  having  groups  of  spines  around  the  dorsal  pore 
which  do  not  reach  the  supradorsal  membrane.  In  other  characters 
much  like  Pteraster.  Numerous  young  were  found  in  the  nidamental 
cavity  by  Professor  Ludwig. 

Professor  Fisher  thinks  that  the  type  was  merely  an  abnormal 
Pteraster  militaris.  Ludwig  mentions  young  taken  from  the  nida- 
mental cavity,  but  does  not  say  that  they  disagreed  with  the  mother, 
as  would  most  likely  have  been  the  case  if  the  absence  of  the  osculum 
was  abnormal. 


278  VERRILL 

PTERASTERIDES  APORUS  (Ludwig)  Verrill. 

Pteraster  aporus  LUDWIG,  Zool.  Jahrb.,  p.  293. 

Pterasterides  aporus  VERRILL,  Amer.  Naturalist,  XLIII,  p.  547,  1909. 

I  have  not  seen  this  species.  The  original  description  follows : 
"  The  radius  of  the  disk  is  35  mm.,  that  of  the  arm,  98  mm. 
Where  the  arms  branch  off  from  the  disk,  the  width  is  from  40  mm. 
to  42  mm.  Both  disk  and  arms  are  somewhat  flattened.  The  supra- 
dorsal  membrane  is  rather  thick,  soft,  with  corrugated  (wrinkled) 
upper  surface,  through  which  calcareous  spines  in  no  wise  protrude ; 
in  its  outermost  layer  are  found,  on  microscopic  investigation, 
numerous  reticulate  little  calcareous  bodies,  from  the  outer  surface 
of  which  small  pointed  spinelets  project.  The  paxillae,  which  support 
the  supradorsal  membrane,  consist  of  a  pedicel  (stalk)  from  2  mm. 
to  3  mm.  long,  from  the  upper  end  of  which,  as  a  rule,  four  much 
thinner  spines,  about  5  mm.  long,  diverge.  On  the  inner  dorsal  skin 
the  paxillae  are  inserted  at  intervals  of  3  mm.,  on  the  average.  The 
calcareous  net  of  the  inner  dorsal  skin,  on  the  knotted  points  of 
which  the  paxillae  stand,  is  made  up  of  more  or  less  elongated  calca- 
reous plates,  which  partly  overlap  the  knotted  points  of  the  network, 
like  tiles.  In  the  circumference  next  to  the  large  anal  opening  into 
the  nidamental  cavity  are  some  noteworthy  thick  paxillae,  which  bear 
on  their  upper  ends  a  great  number  of  spines.  On  the  under  side  of 
these  starfishes  one  may  count,  on  each  arm,  over  fifty  pairs  of  little 
feet  and  a  corresponding  number  of  fins  (or  combs).  Towards  the 
exterior  the  overlying  marginal  border  is  so  wide  that  it  reaches  to 
the  edge  of  the  arms.  All  the  spines  of  the  marginal  border  and  of 
the  combs  are  enveloped  for  their  length  in  a  rather  thick  and  soft 
membrane.  In  each  comb  lie  from  eight  to  nine  spines,  of  which  the 
two  innermost,  standing  next  to  the  ambulacral  furrow,  are  very 
weak  and  much  shorter  than  the  six  to  seven  outer  ones,  which  are 
strong  and  of  about  equal  size.  Between  the  combs  may  be  seen  the 
large  segmental  apertures  furnished  with  clapper-shaped  aperture- 
papillae.  At  the  mouth,  each  jaw  carries  an  oral  comb,  which  bears 
six  spines  ('  mouth-spines  proper,'  Sladen)  and  is  not  united  with  its 
neighbor.  Towards  the  exterior  from  this  comb,  on  each  jaw,  is  a 
light,  curved,  rarely  strong,  single  spine  ('  secondary  mouth-spine,' 
Sladen)  which  likewise  is  covered  by  the  soft  skin. 

"  From  all  the  species  of  Pteraster  now  known  (militaris  Miiller 
and  Troschel,  pulvillus  M.  Sars,  multipes  M.  Sars,  capensis  Gray, 
cribrosus  v.  Mart.,  dance  Verrill,  affinis  Smith,  rugatus  Sladen, 
stellifer  Sladen,  semireticulatus  Sladen,  carribceus  Perrier),  the 


SHALLOW- WATER   STARFISHES  2/9 

present  unique  example,  of  which  unfortunately  no  previous  original 
account  exists,  may  be  distinguished  in  a  conspicuous  manner  by  the 
absence  of  the  so-called  oscular  orifice  of  Sladen,  in  the  supradorsal 
membrane  (comp.  because  of  this  and  some  others  among  the  fol- 
lowing required  characters :  Sladen,  Asteroidea  of  the  '  Challenger ' 
Expedition,  Preliminary  Notice,  Part  I,  Pterasteridae ;  in  Journ. 
Linn.  Soc.,  London,  xvi,  p.  191,  1882).  If  all  other  characters  did 
not  correspond  with  those  of  the  genus  Pteraster,  one  would  be  war- 
ranted in  founding  a  new  genus  on  the  above-mentioned  absence  of 
the  so-called  oscular  orifice.  As  I  propose  to  give  in  another  place  a 
more  detailed  discussion  of  the  organization  of  this  new  species,  and 
a  description  of  the  numerous  quantity  of  young  forms  found  in  the 
nidamental  cavity,  I  shall  content  myself  for  the  present  with  a  short 
account,  setting  forth  the  evident  differences  between  this  and  other 
known  species."  (Ludwig.) 

Bering  Sea,  off  St.  George  Island,  in  30  meters  (Ludwig). 

Genus  Diplopteraster  Verrill. 

Diplopteraster  VERRILL,  Amer.  Journ.  Sci.,  xx,  p.  400,  1880.     Fisher,  op.  cit., 
191 1&,  p.  370. 

Thick,  pentagonal,  or  short-rayed,  with  a  thick  outer  integument 
containing  strong  reticulated  muscle  bands.  Marginal  membranous 
fold  wide  and  thick.  Oral  spines  webbed  together.  Tube-feet  in 
four  rows.  Fans  of  adambulacral  spines  and  furrow-series  alter- 
nately larger  and  smaller,  and  alternately  placed  farther  forward 
and  back. 

In  life,  the  surface  secretes  a  large  amount  of  mucus,  when  taken 
from  the  sea,  and  emits  strong  phosphorescent  light. 

When  fresh  the  surface  appears  smooth  or  nearly  so.  In  pre- 
served and  contracted  specimens  the  larger  paxillary  spines  push  up 
against  or  through  the  dorsal  membrane,  so  as  to  give  it  a  more  or 
less  bristling  or  spinose  appearance,  as  happens  in  most  other 
species  of  Pterasteridae. 

DIPLOPTERASTER  MULTIPES  (Sars)  Verrill. 

Pteraster  multipes  M.  SARS,  Vidensk.  Selskabs.  Forhandl.,  1865,  p.  200;  Fauna 

Litt.  Norvegiae,  p.  65,  pi.  vni,  figs.  1-17,  1877. 
Retaster  ?  multipes  SLADEN,  op.  cit.,  pp.  477,  478,  800,  1889. 
Diplopteraster  mulitpes  VERRILL,  Amer.  Journ.  Sci.,  xx,  p.  400,  1880;  Ann. 

Report  U.  S.  Fish  Comm.,  for  1882,  x,  p.  659,  1884;  op.  cit,  xi,  Expl. 

by  the  Albatross  in  1883,  p.  542,  pi.  14,  fig.  43,  1885;  Distrib.  Echinod., 

op.  cit.,  1895,  p.  202.    Fisher,  op.  cit.,  191  ib,  p.  371,  pi.  cvii,  figs.,  i,  2. 


280  VERRILL 

This  large  North  Atlantic  and  Arctic  species  has  been  recorded 
from  the  Pacific  side  by  Professor  Fisher,  from  several  stations  in 
Bering  Sea  and  off  the  Aleutian  Islands,  in  81  to  350  fathoms,  and 
also  from  off  San  Diego,  California,  in  628  to  640  fathoms. 

On  the  Atlantic  side  it  has  been  found  at  many  stations  between 
N.  lat.  44°  26'  and  37°  07'  50",  in  70  to  640  fathoms.  Most  common 
between  100  and  300  fathoms. 

It  extends  southward  on  the  European  coast  to  Norway.  Also 
recorded  from  Barents  Sea  and  off  Japan.  I  have  seen  no  Pacific 
specimens. 

Although  it  has  not  been  taken  quite  within  the  limits  of  this 
report,  it  is  introduced  here  on  account  of  its  distribution  as  a  cir- 
cumpolar  species. 

When  first  taken  from  the  sea  I  have  observed  that  this  species 
is  highly  phosphorescent.  It  discharges  a  large  amount  of  thick 
mucus  to  which  the  phosphorescence  seems  to  be  due. 

The  integument  and  wide  marginal  fringe,  when  fresh,  are  very 
soft  and  thick. 

Order  PHANEROZONA  Sladen  (emended). 

Phanerosona  (emended)  SLADEN,  Voyage  Challenger,  Zool.,  xxx,  Introduc- 
tion, pp.  xxvni,  XLI,  1889.  Verrill,  Revision,  Trans.  Conn.  Acad.,  x, 
p.  200,  1899.  Fisher,  op.  cit,  191  ib,  p.  17. 

Valvatce  +  Paxillosae  (as  orders)  PERKIER,  in  Mem.  Etoiles  de  mer,  Antilles 
etc.,  p.  154,  1884. 

Valvata  -f-  Paxillosa  PERRIER,  Exped.  Trav.  et  Talism.,  pp.  22,  23,  28,  29,  1894 ; 
Resultats  Camp.  Scientif.  Albert  I,  Prince  de  Monaco,  fasc.  xi,  p.  16,  1806. 

This  order  includes  starfishes  varying  in  form  from  pentagonal 
to  stellate,  with  long  rays.  They  have  two  rows  of  large  and  thick 
marginal  plates  (rarely  but  one  distinct  row,  as  in  Luidia),  which 
are  in  contact  and  usually  form  a  thick,  nearly  perpendicular  margin. 
They  are  nearly  always  larger  than  the  dorsal  and  actinal  plates. 
Papular  pores  are  generally  confined  to  the  upper  surface  and  usually 
placed  singly,  often  in  special  areas.  Dorsal  plates  various.  Their 
surfaces  may  be  closely  covered  with  angular  tesselated  plates,  which 
may  be  naked,  granulated,  or  covered  with  a  smooth  skin;  or  they 
may  take  the  form  of  pseudopaxillae,  protopaxillae,  or  true  paxillae,1 
especially  in  Paxillosa. 

*The  various  forms  of  paxilliform  plates  were  thus  distinguished  by  me 
in  1894: 

"  True  paxillcc  are  columnar  or  hour-glass-shaped  ossicles,  usually  with 
isolated,  circular  bases,  which  bear  at  summit  a  group  of  small  spinules, 
of  which  the  marginal  series  are  usually  different  from  the  rest  and  divergent, 
so  as  to  cover  the  intervening  spaces  between  the  spines,  thus  forming 


SHALLOW- WATER   STARFISHES  28 1 

In  some  families  the  plates  are  reticulated  ossicles,  smooth,  or 
bearing  large  spines  or  tubercles  as  in  Oreasteridse.  In  other  cases 
(Porania,  etc.)  a  thick  skin  may  cover  and  obscure  the  plates. 

The  interactinal  plates  may  be  few  in  stellate  forms  or  many  in  the 
pentagonal  forms.  In  the  latter  cases  they  are  usually  arranged  in 
definite  rows,  most  often  parallel  with  the  adambulacral  plates,  the 
latest  formed  ones  being  situated  next  the  median,  interradial,  mar- 
ginal plates. 

The  interactinal  plates  are  most  commonly  tesselated  and  granu- 
lated, or  else  in  the  form  of  pseudopaxillae  or  protopaxillae ;  but  they 
are  sometimes  spinose.  They  may  be  more  or  less  imbricated  by  their 
edges  or  lobes. 

Adambulacral  plates  are  not  compressed,  rather  large,  usually 
quadrangular,  and  commonly  bear  numerous  small  spines,  various  in 
arrangement. 

Ambulacral  plates  are  rather  large,  not  compressed  nor  crowded. 
Ambulacral  feet  are  in  two  rows,  with  terminal  suckers  in  Valvu- 
losa ;  but  in  the  Paxillosa  large  and  pointed,  or  without  suckers. 

A  median,  dorsal,  nephridial  pore  or  pseudanus  is  generally  pres- 
ent, but  often  absent.  Madreporic  plate  various.  Dentary  or  oral 
plates  are  often  large,  with  numerous  oral  spines. 

fascicles.  These  are  highly  developed  in  most  species  of  Astropecten  and 
Luidia. 

"  Spinopaxilla  are  of  the  same  general  structure,  but  the  center  of  the 
summit  is  occupied  by  a  distinct  spine,  or  by  more  than  one.  Such  forms 
occur  on  Luidia,  Pontaster,  etc. 

"  Parapaxilla  are  lower  and  broader,  rounded,  or  stellate  ossicles,  or 
angular  plates,  with  a  raised  central  portion,  tabulate,  truncate,  or  like  a  low 
cone.  They  may  be  either  isolated  or  articulated  by  their  bases.  The  sum- 
mit is  covered  with  small,  short,  differentiated  spinules,  like  those  of  true 
paxillse.  Those  on  the  dorsal  surface  of  Plutonaster  are  examples.  They 
sometimes  bear  a  central  spine. 

"Protopaxillae  are  similar,  but  less  elevated,  convex  ossicles  or  plates, 
covered  with  round  or  angular  granules,  with  the  marginal  series  dif- 
ferentiated and  more  or  less  covering  the  grooves  between  the  plates.  As 
in  the  preceding,  there  may  be  a  central  spine  in  some  cases.  This  form  also 
occurs  on  Plutonaster,  and  on  many  species  of  Goniasteridae.  The  transition 
from  this  last  kind  to  simple,  uniformly  granulated  plates  is  easy,  when  the 
grooves  between  the  plates  become  obsolete. 

"  Pseudopaxillce  are  articulated  plates  with  a  flattened,  usually  lobed,  and 
often  overlapping  base,  which  bear  a  group  of  slender,  fascicled  or  divergent 
spinules  on  the  more  or  less  raised  central  or  subcentral  area  or  boss.  These 
have  no  differentiated  marginal  series  of  spinules.  This  form  is  seen  in 
Solaster,  Henricia,  etc." 


282  VERRILL 

Pedicellariae  are  sometimes  lacking,  but  usually  present  and  some- 
times large.  They  may  be  bivalvular,  sessile  and  seated  over  a  pore 
(foraminate),  or  implanted  in  special  pits  on  the  plates  (fossate)  ;  or 
else  papilliform  or  fasciculate,  composed  of  two  to  four  or  more 
modified,  convergent  spinules ;  or  pectinate,  consisting  of  two  comb- 
like  groups  of  spinules  convergent  over  a  suture  between  two 
adjacent  plates,  as  in  Luidiaster  (see  pi.  xxxiv). 

The  sessile  valvular  pedicellariae  with  a  pore  between  the  valves 
are  called  foraminate;  they  may  have  two,  or  more  than  two,  valves 
(bivalvular  and  trivalvular,  etc.).  In  the  suborder  Valvulosa  they 
are  often  provided  with  a  pair  of  pits  or  fossae,  into  which  the  valves 
fit  when  widely  open.  The  valves  in  these  are  often  spatulate,  spoon- 
shaped,  tongs-shaped,  or  battledore-shaped,  and  usually  may  best 
be  called  plataleiform  or  spatulate.  When  furnished  with  receiving 
pits,  they  have  been  called  excavate;  but  fossate  seems  to  be  a  more 
appropriate  term. 

The  peculiar  papilliform  pedicellariae  of  the  Paxillosa,  with  two  to 
four  or  more  slender  valves,  apparently  formed  by  modified  spinules 
or  granules,  are  also  found  in  certain  of  the  Valvulosa.  Sometimes 
such  pedicellariae  are  found  associated  with  larger  valvular  pedicel- 
lariae on  the  same  specimen,  as  in  the  genus  Nymphaster,  and  in  a 
few  other  genera. 

The  writer,  in  a  revision  of  the  classification  of  the  Asterioidea, 
in  1899,  proposed  a  rearrangement  of  the  families  and  subfamilies. 
That  general  arrangement  is  here  adopted,  but  with  various  changes 
made  necessary  by  subsequent  discoveries.1 

Order  PHANEROZONA. 
Suborder  I.  VALVULOSA  Verrill  =  VALVATA  Perrier 

(sense  extended). 
Family      I.  OPHIDIASTERIDM   Verrill,    1867 = Lmckiada 

Perrier,  1875. 

Family    II.  VALVASTERIDM  Fisher,  1911. 
Family  III.  ASTEROPID2E  Fisher,  1911. 
Family  IV.  OREASTERW2E   Fisher  =  Pentaterotida   Gray 

(restricted). 

Family     V.  M1MASTERID2E  Verrill. 

Family   VI.  GONIASTERIDJE  Forbes  (restricted),  includes 
Antheneida  Perrier  (restricted). 

1Dr.  Fisher  (op.  cit.,  191  ib)  has  adopted  a  very  similar  arrangement  of  the 
families  and  subfamilies,  with  several  changes,  which  are  here  mostly  ac- 
cepted. Some  small  extralimital  groups  are  not  included  in  this  table. 


SHALLOW-WATER   STARFISHES  283 

Subfamily         I.  GONIASTERINJE  Verrill  =  Pentagonas- 

terina  Sladen  (pars). 

Subfamily       II.  PSEUDARCHASTERINJE  Sladen. 
Subfamily      III.  MEDIASTERIN2E  Verrill,  1899. 
Subfamily      IV.  NECTRIINJE  Perrier;  Fisher. 
Subfamily       V.  HIPPASTERIINJE  Verrill. 
Subfamily      VI.  ANTHENEIN2E  Fisher  (sense  restricted). 
Subfamily    VII.  LEPTO  GO  MASTERING  Perrier. 
Subfamily  VIII.  CHITON ASTERINJE  Fisher. 

Family  VII.  ODONTASTERIDJE  Verrill  =  Gnatkosteriwe 
Perrier  (pars). 

Family  VIII.  CH&TASTERID1E  Ludwig. 

Family  IX.  ARCHASTERID2E  Verrill  (restricted  to  Arch- 
aster). 

Suborder  II.  MYONOTA  Ludwig. 

Family  X.  BENTHOPECTINIDJE  Verrill  =  Benthopec- 
tinince  Verrill,  1894. 

Subfamily    I.  BENTHOPECTININJE  Verrill,  1894. 
Subfamily  II.  PONTASTERIN&.  Verrill,  1899. 

Suborder  III.  PAXILLOSA  Perrier  (sense  restricted). 
Family      XI.  PORCELLANASTERWM  Sladen. 

Subfamily    I.  PORCELLASTERINM  Sladen. 
Subfamily  II.  CTENODISCINJE  Sladen. 

Family    XII.  GONIOPECTINIDJE  Verrill. 

Family  XIII.  ASTROPECTINIDJE  Gray  (restricted). 

Family  XIV.  LUIDIID^E  Verrill  =  Luidiina  Sladen. 

It  will  be  noticed  that  in  the  above  arrangement  the  family  Arch- 
asteridias  of  Perrier,  1894,  is  divided  among  five  distinct  families 
and  subfamilies.  Most  of  his  genera  are  now  placed  in  the  Pseud- 
archasterinse. 

The  subordinal  name  Valvata  I  have  here  changed  to  Valvulosa, 
partly  because  it  is  not  strictly  identical  with  Valvata  of  Perrier, 
and  partly  to  make  its  termination  correspond  with  Paxillosa. 

In  the  arrangements  of  Perrier,  and  of  Sladen,  Valvaster  is  made 
a  subfamily,  Valvasterinae,  of  the  family  Echinasteridae.  Fisher 


284  VERRILL 

makes  it  a  family  of  Spinulosa.  The  reasons  for  this  are  not  very 
evident,  for  Valvaster  has  a  series  of  large,  broad,  valvular  pedicel- 
laricE  on  the  supermarginal  plates,  which  are  not  otherwise  found 
in  the  family  Echinasteridae,  nor  in  the  order  Spinulosa.  Moreover, 
it  has  erect  bivalve  pedicellariae  on  the  actinal  and  adambulacral 
plates,  with  large  marginal  plates  in  two  rows,  and  is,  therefore, 
phanerozonate.  These  characters  and  its  jaws  and  odontophore 
indicate  that  it  belongs  to  the  suborder  Valvulosa. 

It  should,  in  my  opinion,  form  a  separate  family,  Valvasteridae, 
near  Ophidiasteridae,  but  peculiar  in  having  the  dorsal  ossicles  reticu- 
lated and  bearing  solitary  spines  in  longitudinal  rows.  Odontophore 
with  articulating  tubercles,  as  in  Ophidiasteridae;  jaws  pointed. 
Actinal  plates  bear  flattened  spines.  The  ambulacral  suckers  have  a 
rosette  of  calcareous  deposits. 

The  former  subfamily  Mimasterinae  I  have  here  raised  to  the  rank 
of  a  family,  to  include  only  the  genus  Mimaster,  which  seems  to  be 
a  rather  isolated  form. 

The  families  of  the  above  list  represented  within  our  limits  are 
Nos.  I,  III,  VI,  VII,  X,  XI,  XIII,  XIV. 

Suborder  VALVULOSA  Verrill  =  VALVATA  (Perrier), 

emended. 

Valvulata  (order)  PERKIER,  Nouv.  Arch.  Mus.  Hist  Nat.,  vi,  1885. 
Valvata  (order)  PERRIER,  Exped.,  Trav.  et  Talism.,  pp.  22,  23,  28,  29,  1894. 
Valvata  (suborder)   VERRILL,  Revision,  Trans.  Conn.  Acad.,  x,  p.  200,  1899. 

This  extensive  group  includes  starfishes  with  stellate  or  pentagonal 
forms,  usually  rather  stiff  in  appearance,  and  nearly  always  five- 
rayed,  in  which  the  dorsal  plates  are  most  commonly  tesselated  and 
granulated,  or  else  take  on  the  character  of  protopaxillae  or  pseudo- 
paxillae.  Sometimes  they  are  naked ;  often  spinose ;  very  rarely  truly 
paxilliform;  sometimes  covered  by  a  thick,  smooth,  or  granulated 
integument. 

The  various  kinds  of  abactinal  ossicles  pass  into  each  other  by 
various  intermediate  forms,  so  that  it  is  impossible  to  draw  any  very 
strong  or  sharp  family  lines  on  this  character  alone,  though  the 
character  of  the  plating  may  generally  be  taken  as  of  generic  value. 
Sometimes  they  are  openly  reticulate;  rarely  (Asteropidae)  they  are 
partly  abortive. 

Pedicellariae  are  often  lacking;  when  present  they  are  usually 
valvular,  foraminate  and  sessile,  or  fossate.  The  valves  are  either 
granuliform,  or  large  and  broad,  or  narrow  and  spoon-shaped, 
plataleiform,  spatulate,  or  battledore-shaped,  rarely  papilliform. 


SHALLOW-WATER   STARFISHES  285 

Actinal  plates  are  usually  rather  numerous  and  tesselated  or 
imbricated.  Adambulacral  spines  are  usually  numerous  and  papilli- 
form  or  spiniform.  A  distinct  dorsal  pore  (pseudanus)  is  usually 
present.  Genital  pores  generally  ten,  usually  all  separate,  situated 
in  the  dorsal  interradial  areas ;  sometimes  serial  on  the  sides  of  the 
rays.  Oral  plates  usually  large  and  convex  on  the  actinal  surface. 

Superambulacral  plates  are  present  in  some  genera,  absent  in 
others.  The  existence  of  superambulacral  plates  has  formerly  been 
supposed  to  be  characteristic  of  the  Astropectinidae  only,  but  they 
exist  in  several  of  the  genera  here  referred  to  Valvulosa  (viz., 
Mediaster,  Dy taster,  Pseudarchaster,  etc.),  and  in  other  groups. 

Adambulacral  plates  wide,  with  a  longitudinal  group  of  small 
furrow-spines  and  with  one  or  several  spines  on  the  actinal  side, 
often  becoming  much  longer  distally.  Dentary  or  jaw-plates  large 
and  usually  convex  on  the  actinal  surface,  bearing  numerous  oral  and 
epioral  spines. 

Ambulacral  feet  large,  with  suckers.  Owing  to  the  generally  close 
union  of  the  ossicles,  especially  of  the  large  marginal  plates,  most 
of  these  starfishes  are  rather  rigid,  showing  much  less  flexibility  than 
in  most  other  groups.  The  Ophidiasteridae  are  exceptional  in  this 
respect,  for  they  are  mostly  very  flexible. 

Family  GONIASTERIDJE  (Forbes),  emended  Verrill. 

Goniasterida  (pars)  FORBES,  1840.    Verrill,  Trans.  Conn.  Acad.  Sci.,  i,  p.  343, 

1867.     Perrier  (pars),  Revision,  Arch.  Zool.  Exper.  et  Gen.,  iv,  pp.  281, 

283,  289,  291,  1875 ;  op.  cit,  v,  p.  I,  1876. 
Pentagonasterina  VIGUIER  (pars),  subfamily,  op.  cit,  vn,  p.  166,  pi.  x,  figs. 

20-25,  1878. 

Pentacerotidce  (pars)  GRAY,  Synopsis,  p.  5,  1866. 
Pentagonasterida  (pars)   PERRIER,  op.  cit.,  p.  231,  1884.     Sladen  (pars),  op. 

cit.,  pp.  260,  264,  1889. 
Goniasteridce  VERRILL  (restricted),  Revision,  in  Trans.  Conn.  Acad.  Sci.,  x, 

p.  145,  1899  (non  Viguier).    Fisher,  op.  cit.,  191  ib,  p.  158. 

Phanerozonate  starfishes  usually  having  a  rather  broad,  flat  or 
slightly  convex,  rather  rigid  disk,  sometimes  nearly  pentagonal  in 
form,  but  often  stellate  with  more  or  less  prolonged  rays.  Mar- 
ginal plates  usually  large  and  thick,  forming  a  thick,  nearly  vertical 
margin,  the  two  rows  equal  or  subequal. 

Dorsal  plates  various,  but  usually  tesselated,  polygonal,  or  round- 
ish, sometimes  lobed  or  substellate,  or  united  by  internal  ossicles. 
They  are  commonly  granulated  or  protopaxilliform,  sometimes 
spinulose,  or  bear  tubercles  or  spines ;  rarely  naked,  or  covered  with 
soft  skin,  with  or  without  granules  or  pedicellariae. 


286  VERRILL 

Interactinal  plates  usually  numerous,  angular,  tesselated,  or  imbri- 
cated. Superambulacral  plates  may  be  present  or  absent.  Tube-feet 
are  in  two  rows  and  have  suckers. 

Pedicellariae  usually  present,  often  large,  usually  bivalve,  but  may 
be  multivalved,  foraminate,  more  often  f ossate.  They  may  occur  on 
any  of  the  plates,  or  on  the  thick  skin  that  covers  them  in  some 
genera  (Anthenea,  etc.). 

NOTE  ON  THE  EARLY  GENERIC  NOMENCLATURE  OF  THE 
GONIASTERID^E.1 

The  name  Pentagonaster,  as  used  by  Linck,  designated  a  composite 
group  of  pentagonal  starfishes  of  this  family  and  others ;  but  Linck 
was  not  a  binomial  writer.  Gray,  1840,  was  the  first  binomial  writer 
to  adopt  Linck's  name.  He  restricted  it  to  a  small  group,  afterwards 
named  Stephanaster  by  Ay  res  (type  P.  pulchellus) ,  chiefly  distin- 
guished by  the  swollen  distal  marginal  plates.  To  this  type  Pentag- 
onaster must  evidently  be  restricted. 

Goniaster  L.  Agassiz,  1835.*  This  generic  name  was  also  intended 
to  include  all  the  pentagonal  starfishes  then  known  to  him,  that  are 
now  referred  to  Goniasteridae  and  Oreasteridae. 

Among  the  species  named  by  Agassiz  was  G.  tesselatus  Lam. 
J.  E.  Gray,  in  1840,  restricted  Goniaster  to  the  latter,  as  he  had  a 
perfect  right  to  do,  and  his  use  of  the  name  must  be  maintained. 
For  another  of  the  species  (equestris),  cited  by  Agassiz,  Gray 
established  the  genus  Hippasteria,  at  the  same  time.  The  first  and 
the  third  species  were  placed  by  Gray  in  his  genus  Pentaceros,  now 
Oreaster  (M.  andTr.). 

The  species  named  by  Agassiz  (see  foot  note  2,  this  page)  thus  be- 
came the  types  of  three  genera  in  Gray's  system.  There  is  no  reason, 
under  the  accepted  rules  of  priority,  for  changing  his  application  of 
the  first  two  of  these  generic  names,  for  his  types  are  all  well- 
known  species. 

It  is  possible,  however,  that  it  may  eventually  be  thought  by  some 
desirable  to  unite  Gray's  three  genera — Goniaster  (restr.),  Pentag- 

1  See  also  Verrill,  Revision  of  Certain  Genera  and  Species  of  Starfishes, 
etc.,  Trans.  Conn.  Acad.  Sci.,  x,  pp.  146-162  (Goniasteridae).  Fisher,  W.  K., 
Necessary  Changes  in  the  Nomenclature  of  Starfishes,  Smithsonian  Miscell. 
Coll.,  LII,  1908,  pp.  91-93  (historical),  and  op.  cit,  19116,  p.  163. 

*The  species  of  Goniaster  cited  by  Agassiz  (Prodromus,  1835,  p.  191)  are 
as  follows :  First,  G.  reticulatus  (now  Oreaster)  ;  second,  G.  equestris  (now 
Hippasteria  phrygiana*)  ;  third,  G.  nodosus  (now  Oreaster)  ;  fourth,  G.  tessela- 
tus  (the  type  of  Goniaster  Gray). 


SHALLOW-WATER   STARFISHES  287 

onaster  (restr.),  and  Tosia  (restr.) — into  one  large  genus,  for  they 
are  closely  related.  In  that  case  the  name  Goniaster  should  be 
applied  to  the  entire  group,  under  which  the  other  two  names  might 
be  retained  for  subgenera  or  sections.  Goniaster  is  certainly  the 
earliest  of  these  names,  under  the  binomial  system. 

The  restriction  of  Goniaster  to  a  species  not  mentioned  by  Agassiz 
(obtusangulus  Lam.),  as  has  been  done  by  Perrier  and  by  Sladen, 
is  entirely  contrary  to  the  rules  of  nomenclature,  and  quite  unjustifi- 
able. Therefore  the  latter  was  made  the  type  of  a  genus  (Pseudore- 
aster)  by  me  in  1899. 

Astrogonium  of  Miiller  and  Troschel,  1842,  should  properly  have 
been  dropped  from  the  system  long  ago.  It  was  a  very  heterogeneous 
group,  composed  (as  the  authors  themselves  stated),  of  several  of 
Gray's  genera  recombined.  This  unnatural  combination  has  been 
rejected  by  many  subsequent  writers.  It  is  a  synonym  of  Gray's 
genera. 

Astrogonium  was  nearly  equivalent  to  the  typical  Goniasterinae, 
collectively. 

Hippasteria.  The  type  of  the  latter  was  the  first-named  species 
of  Astrogonium. 

Gray,  1866,  endeavored  to  restrict  Astrogonium  to  a  minor  group, 
but  his  first  and  most  typical  species  (granularis)  is  a  Tosia,  accord- 
ing to  Gray's  definition  of  the  latter.  Others  belong  to  very  different 
genera. 

Sladen  restricted  it  to  the  group  named  Pentagonaster  by  Gray 
and  Stephanaster  by  Ayres,  which  is  contrary  to  the  generally 
accepted  rules  of  priority.  Other  attempts  to  restore  it  have  been 
equally  unfortunate.  Perrier  formerly  used  it  for  a  section  of 
Pentagonaster. 

Goniodiscus  Miiller  and  Troschel  embraced  several  diverse  genera 
of  Gray's  system  (as  stated  by  them),  such  as  Paulia,  Anthenea, 
Randasia,  Nectria,  Hosea.  Their  first  species  (pentagonulus) ,  pre- 
sumably intended  as  the  type,  was  also  the  type  of  Gray's  Anthenea. 
Two  of  their  species  belong  to  Tosia  Gray. 

Perrier,  and  also  Sladen,  have  restricted  it  to  several  species 
referred  to  Hosea  by  Gray  in  1847  and  1866.  The  latter  genus  was, 
however,  founded  by  Gray  in  1840,  with  a  single  species  (H.  naves- 
cens  Gray)  as  the  type.  Perrier,  who  has  studied  this  type,  states 
that  it  is  a  true  Anthenea.  If  this  be  so,  Hosea  becomes  a  synonym 
of  the  latter. 


288  VERRILL 

Gray,  in  his  later  works,  referred  to  Hosea  several  additional 
species,  which  were  described  under  Goniodiscus  by  Miiller  and 
Troschel. 

The  restriction  of  Goniodiscus  to  this  residual  group  of  species  by 
Perrier,  1875,  was  indefensible  as  well  as  undesirable.  This  group 
of  species/  as  thus  restricted,  was  characterized  by  having  the 
abactinal  radial  plates  "  stellate,"  or  polygonal  with  notched  sides, 
and  by  small,  narrow,  forceps-like  pedicellariae,  usually  scarcely 
larger  than  the  granules,  and  much  like  those  of  Tosia. 

But  if  the  type  is  a  young  Culcita,  as  stated  by  Clark,  the  name  is 
obsolete.  At  least  one  species  (Anthenea  granulifera  Gray),  re- 
ferred to  Goniodiscus  by  Perrier,  who  described  the  figured  type, 
should  probably  be  generically  separated,  for  it  has  short  valvular 
pedicellariae. 

The  above  discussion  was  written  almost  exactly  in  its  present 
form  in  1904,  and  was  an  extension  of  the  notes  on  the  same  subject 
published  by  the  writer  in  1899.  (Revision  of  Genera  and  Species 
of  Starfishes.) 

Professor  Fisher  (19116,  pp.  163-169)  has  since  discussed  the 
same  subject  in  much  the  same  way.  His  conclusions  are  so  closely 
in  accord  with  my  own  that  it  might  seem  superfluous  to  print  my 
results  here.  It  is,  perhaps,  useful  to  do  so,  on  the  principle  that  the 
testimony  of  two  is  better  than  that  of  one.  In  this  case,  also,  Dr. 
Fisher  has  been  able  to  study  the  same  types  of  Perrier  that  I  had 
previously  studied,  in  the  Museum  of  Comparative  Zoology,  which 
adds  to  the  value  of  the  results. 

Another  good  reason  for  repeating  these  discussions  is  the  fact 
that  the  prestige  of  the  great  work  of  Sladen  and  the  several  impor- 
tant works  of  Perrier  has  caused  the  adoption  of  untenable  nomen- 
clature, even  up  to  the  present  time,  especially  by  several  European 
writers. 

In  a  very  recent  work  (op.  cit.,  1912)  Ludwig  has  very  fully  and 
satisfactorily  discussed  the  genera  Pentagonaster  Gray  and  Tosia 
Gray,  retained  in  the  same  sense  as  by  me,  in  1899,  and  by  Fisher  in 
1911. 

1This  group  has  recently  (1906)  been  renamed  Goniodiscides  by  Fisher. 
Type  G.  sebce  (Miiller  and  Troschel),  non  Gray.  He  properly  rejected 
Goniodiscus,  because  it  was  essentially  a  synonym  of  Gray's  genera.  In  my 
remarks  on  this  group  (1899,  p.  149),  the  type  was  inadvertently  stated  as 
cuspidatus,  instead  of  sebce.  But  Dr.  H.  L.  Clark  has  shown  that  seba  is 
only  a  young  Culcita. 


SHALLOW- WATER   STARFISHES  289 

He  recognizes  four  species  in  each  genus,  thus  reducing  many 
nominal  species  to  synonyms. 

These  genera  are  not  found  on  either  coast  of  America.  The 
species  are  mostly  Australian. 

Subfamily  GONIASTERIN&  Verrill. 

Goniasterince  VERRILL,  Revision,  p.  145,  1899.     Fisher,  op.  cit.,  191  ib,  p.  196, 

(includes  Mediasterince). 
Pentagonasterince  (pars)  SLADEN,  Voy.  Challenger,  xxx,  pp.  262,  264,  1889. 

In  this  extensive  group  the  dorsal  and  actinal  plates  are  generally 
tesselated,  polygonal,  or  rounded,  often  stellate,  in  pretty  close  con- 
tact, or  else  with  intervening  plates  or  ossicles  of  small  size.  They 
may  be  either  granulated,  or  else  smooth  and  naked,  except  for  a 
marginal  series  of  granules ;  sometimes  they  bear  large  conical  tuber- 
cles or  spines.  Papulae  usually  placed  singly  around  the  radial  plates. 
Pedicellariae,  when  present,  foraminate  and  usually  small,  commonly 
with  slender,  flat,  spoon-shaped  or  spatulate  valves  (plataleiform) 
and  f ossate ;  sometimes  with  large,  wide,  plain,  short  valves. 

Genus  CERAMASTER  Verrill  (as  subgenus),  Fisher. 

Astrogonium  (pars)  MULLER  and  TROSCHEL,  Syst.  Ast.,  1842. 
Pentagonaster  (pars)  PERKIER,  Revision,  p.  20,  1876  (non  Gray,  1840). 
Pentagonaster  (pars)   SLADEN,  Voy.  Chall.,  xxx,  p.  264,  1889  (non  Gray). 

Perrier,  Exp.  Trav.  et  Talism.,  pp.  389,  390,  1894. 
Tosia,  subgenus  Ceramaster  VERRILL,  Revision  Genera,  etc.,  in  Trans.  Conn. 

Acad.,  x,  pp.  148,  158,  160,  1899. 
Ceramaster  FISHER,  op.  cit,  191  ib,  p.  204. 
Philonaster  KCEHLER,  Investigator  Exped.,  1909,  p.  74  (teste  Fisher). 

Form,  short-rayed  stellate.  The  marginal  plates  are  very  regular 
and  generally  correspond  pretty  closely  in  the  upper  and  lower  series, 
except  distally;  an  odd  interradial  plate  sometimes  occurs  abnor- 
mally; they  are  usually  closely  granulated,  but  sometimes  have  a 
small,  naked  central  area,  with  rows  of  granules  around  the  margins. 
Apical  plate  small.  The  abactinal  plates  of  the  radial  areas  are 
granulated,  somewhat  tabulate,  their  bases  most  often  hexagonal  or 
roundish,  with  lobes,  and  crowded  pretty  closely  together,  without 
special  intervening  or  radiating  connective  ossicles,  and  usually 
without  many  secondary  plates  of  small  size ;  they  sometimes  extend 
to  the  apical  plates.  Superambulacral  plates  are  lacking. 

Interactinal  plates  are  tesselated  and  granulated.  The  papular 
pores  are  usually  rather  numerous,  generally  placed  singly  in  the 
angles  between  the  plates  of  the  basal  radial  areas,  and  sometimes 

20 


290  VERRILL 

on  the  central  part  of  the  disk,  but  not  on  the  triangular  interradial 
areas,  where  the  plates  are  angular  and  closely  in  contact. 

The  pedicellariae  are  often  lacking ;  when  present  they  are  small, 
elevated,  usually  with  two,  rarely  three,  spatulate  or  spoon-shaped 
blades,  higher  than  broad,  and  set  in  special  fossae  or  pits  of  similar 
shape,  in  the  surface  of  the  plates.  They  may  occur  on  any  or  all 
kinds  of  plates,  either  above  or  below,  or  on  both  sides. 

The  adambulacral  spines  are  numerous,  small  and  crowded.  They 
grade  more  or  less  into  the  actinal  granulation ;  the  marginal  furrow- 
series  form  a  simple  row  of  two  to  six  or  more ;  they  are  usually  not 
much  longer  than  those  of  the  next  series  and  not  separated  from 
them  by  a  wide  space.  Distally,  some  of  the  spines  of  the  second 
series  usually  become  much  larger  and  longer  than  the  rest. 

CERAMASTER  GRANULARIS  (Retzius)  Verrill. 

Plate  iv,  figures  I,  2;  plate  L,  figures  2,  20  (details). 
Asterias  granularis  RETZIUS,  K.  Vet.  Akad.  Nya  Hand!.,  iv,  p.  238,  1783. 

Abildgaard,  in  Zool.  Dan.,  fas.  in,  p.  19,  pi.  xcn,  1788.    Retzius,  Diss. 

Syst.  Aster.,  p.  10,  1805. 
Astrogonium  granulare  MULLER  and  TROSCHEL,  Syst.  Asteriden,  p.  57,  1842. 

M.  Sars,  Oversigt  af  Norges  Echinodermer,  p.  46,  1861.    Gray,  Synopsis, 

p.  10,  pi.  i,  fig.  4,  1866.    Verrill,  Expl.  by  the  Albatross  in  1883,  p.  542, 

pi.  XVTII,  figs.  48  and  480,  1885.    Whiteaves,  Trans.  Royal  Soc.  Canada,  iv, 

p.  117,  1887. 

Goniaster  granularis  LUTKEN,  Vidensk.  Medd,  nat.  Foren.,  p.  146,  1865. 
Pentagonaster  granularis  PERKIER,  Revis.  Stell.  du  Mus.  Arch,  de  Zool.  Exper. 

et  Gen.,  v,  p.  40  (224),  1876.    Sladen,  Voy.  Challenger,  xxx,  p.  268,  1889. 

Bell,  Catal.  British  Echinod.  in  British  Museum,  p.  73,  pi.  x,  figs.  4,  5,  6, 

1892.    Verrill,  Distrib.  of  Echinod.,  Amer.  Journ.  Sci.,  xux,  p.  135,  1895. 

Danielssen  and  Koren,  Asteroidea,  Norske  Nordhavs  Exped.,  Zool.,  xi, 

p.  58,  1884. 
Tosia  (Ceramaster)  granularis  VERMLL,  Revision  Genera,  etc.,  in  Trans.  Conn. 

Acad.  Sci.,  x,  p.  162,  1899.    Whiteaves,  Catal.  Invert.  E.  Canada,  p.  49, 

1901. 

The  marginal  plates,  above  and  below,  are  closely  covered  with 
similar  and  small  granules.  The  plates  of  the  dorsal  surface  are 
hexagonal  on  the  radial  areas  at  the  bases  of  the  rays,  and  are  mostly 
transversely  elongated,  and  surrounded  by  six  papular  pores,  corre- 
sponding to  the  angles.  In  the  interradial  areas  they  are  transversely 
rhombic,  often  with  the  acute  angles  truncated,  where  pores  inter- 
vene. All  are  closely  covered  with  small  angular  granules. 

The  actinal  interradial  plates  are  crowded,  polygonal,  and  closely 
covered  with  small,  polygonal,  granule-like  spinules  with  rounded 
tips,  about  thirty  on  the  larger  plates,  their  size  decreasing  toward 
the  marginal  plates,  where  they  are  very  small. 


SHALLOW- WATER   STARFISHES  29! 

The  inner  adambulacral  spinules  form  a  simple  marginal  row, 
with  three  or  four  spines  on  each  plate,  of  which  the  proximal  is 
smaller  and  sets  farther  back,  so  as  to  be  partly  overlapped  by  the 
distal  one  of  the  preceding  plate ;  the  others  are  rather  short,  stout, 
blunt,  scarcely  tapered,  about  as  long  as  the  breadth  of  the  adambu- 
lacral plates.  Outside  the  furrow-series,  each  plate  bears  an  actinal 
group  of  about  seven  to  eleven  short,  stout  polygonal  spinules  or 
granules,  one  of  which  occupies  the  center,  while  the  others  surround 
it;  those  on  the  side  next  the  furrow-series  are  much  larger  and 
somewhat  longer  than  the  rest. 

Oral  spinules  numerous,  short,  stout,  polygonal,  seven  or  eight  on 
the  border  of  the  dentary  plate,  and  a  median  or  sutural  epioral 
group,  consisting  of  a  row  of  six  to  eight  on  each  plate,  with  two 
shorter  intermediate  or  central  rows  of  three  or  four  smaller  ones. 

This  is  a  truly  arctic  and  circumpolar  species.  The  description 
above  is  from  a  North  Atlantic  specimen,  collected  at  Albatross 
Station,  No.  2506,  U.  S.  National  Museum  (Catalog  No.  11428).  I 
have  seen  no  well  preserved  West  Coast  specimens  perfect  enough  to 
figure. 

Off  mouth  of  Qualicum  River,  Straits  of  Georgia,  in  40  fathoms 
(coll.  G.  M.  Dawson,  teste  Whiteaves)  ;  Marmot  Island,  Alaska 
(Ives). 

It  has  been  taken  off  the  northeastern  American  coast  at  several 
stations,  between  N.  lat.  44°  28'  30"  and  41°  4/,  by  the  U.  S.  Fish 
Commission.  Often  taken  by  the  Gloucester,  Massachusetts,  fisher- 
men on  the  fishing  banks,  off  Nova  Scotia  and  Newfoundland,  in  50 
to  200  fathoms.  Occurs  off  the  northern  coasts  of  Norway  and 
Great  Britain,  and  in  the  Arctic  Ocean  generally. 

Bathymetrical  range,  50  to  471  fathoms  on  the  east  American  coast, 
rarely  taken  below  150  fathoms. 

Mr.  Ives  recorded  a  specimen  from  Monterey,  California,  which 
is  a  very  unlikely  locality.  Perhaps  his  specimen  was  Mediaster 
equalis. 

CERAMASTER  PATAGONICUS  (Sladen)  Fisher. 
Ptntogonaster  patagonicus  SLADEN,  op.  cit,  1889,  p.  269,  p.  46,  figs.  3,  4;  pL 

XLIX,  figs.  3,  4.     (Off  Patagonia.) 
Ceramaster  patagonicus  FISHER,  op.  cit.,  19116,  p.  214,  pi.  xxxvn,  fig.  4;  pi. 

xxxvm,  figs.  I,  2 ;  pi.  LX,  fig.  3. 

Professor  Fisher  has  described  and  figured  a  species  that  he  thinks 
identical  with  the  one  described  by  Sladen  from  Patagonia,  but 
without  comparison  of  types. 


292  VERRILL 

The  North  Pacific  form  is  a  deep-water  species  and  hardly  comes 
within  my  limits,  though  Fisher  has  one  locality  in  41  fathoms,  and 
one  in  56  fathoms,  off  Unalaska,  in  Bering  Sea.  Other  northern 
localities  recorded  by  him  are  in  68  to  174  fathoms,  from  Bering 
Sea  to  Southeast  Alaska,  Behm  Canal,  4  (?)  to  134  fathoms.  Also 
one  specimen  from  Gulf  of  California,  Carmen  Island. 

Possibly  his  northern  specimens  are  like  those  placed  by  me  under 
C.  granularis.  The  two  species  are,  apparently,  very  similar. 

Dr.  Fisher  also  describes  the  allied  species,  C.  japonicus  (Sladen), 
from  deeper  water,  in  Bering  Sea  and  off  Oregon,  184  to  786  fath- 
oms; and  C.  leptoceramus  ¥.,  from  off  Southern  California,  in  216 
to  638  fathoms  (op.  cit.,  191  ib,  p.  210,  pis.  xxxix,  LVIII,  LX). 

Genus  TOSIASTER  Verrill,  nov. 
Type,  T.  arcticus  Verrill. 

Pentagonal,  with  short  rays.  Marginal  plates  large,  thick, 
coarsely  granulated.  Dorsal  plates  are  stout,  closely  granulated; 
parapaxillae  with  lobed  or  stellate  bases,  and  tabulate  tops.  Papulae 
widely  distributed,  not  confined  to  the  radial  areas,  usually  in  small 
groups  of  two  to  five. 

Adambulacral  plates  are  notably  short;  actinal  side  with  four  to 
six  granuliform  spinules;  furrow-spines  usually  two  or  three  to  a 
plate. 

Interactinal  plates  nearly  rhombic,  closely  covered  with  angular 
granules. 

Bivalve  pedicellariae  occur  on  the  marginal,  interactinal,  and 
adambulacral  plates.  Superomarginal  plates  often  have  a  naked 
central  area. 

TOSIASTER  ARCTICUS  Verrill. 
Plate  L,  figures  3,  30;  plate  xcix,  figures  I,  2  (type). 

Tosia  arctica  VERRILL,  Amer.  Journ.  Sci.,  xxvin,  p.  63,  July,  1909,  figs.  8-8a. 
Ceramaster  arcticus  FISHER,  op.  cit.,  IQII&,  p.  219,  pi.  XL,  figs,  i,  2;  pi.  LVIH, 
fig.  i ;  pi.  LX,  fig.  i. 

The  disk  is  pentagonal  with  short  rays  and  thick,  rounded  mar- 
gins. In  alcohol  the  granules  and  most  of  the  outlines  of  the  dorsal 
plates  are  obscured  by  a  soft,  membranous  or  mucous  structure. 
The  marginal  granules  of  adjacent  plates  are  in  contact  and  cover 
narrow  channels  between  the  plates,  which  have  a  low,  round, 
columnar  form  when  the  granules  are  removed.  Very  small  papulae, 
standing  in  small  groups,  surround  the  plates. 


SHALLOW-WATER   STARFISHES  293 

Radii  of  the  type,  31  mm.  and  45  mm.  to  50  mm.  It  grows  to  a 
much  larger  size. 

The  marginal  plates  are  coarsely  granulated.  The  upper  ones 
often  have  a.  bivalve  pedicellaria  near  the  inner  edge.  The  granules 
are  unequal  in  size,  angular,  closely  in  contact,  about  eight  in  a  trans- 
verse row  on  the  larger  plates.  The  marginal  plates  in  the  interradial 
region  are  squarish  and  pretty  regular,  and  paired  above  and  below ; 
distally  they  become  irregular,  polygonal,  with  rounded  corners,  and 
often  appear  broken ;  they  partially  alternate  in  the  two  rows.  The 
subapical  plates  are  small;  the  apical  or  ocular  plate  is  very  small 
for  this  genus. 

The  lower  marginal  plates  are  closely  granulated  nearly  all  over, 
except  a  very  small  naked  spot  beneath.  The  upper  ones  also  have 
a  large  naked  spot,  near  the  upper  margin,  covering  about  one-fifth 
of  the  surface  of  the  plate,  perhaps  where  the  granules  have  been 
accidentally  rubbed  off.  The  actinal  plates  are  large,  mostly 
squarish ;  about  three  correspond  to  the  breadth  of  each  inferomar- 
ginal  plate,  proximally. 

The  dorsal  plates  are  unequal  in  size,  crowded,  larger  than  in 
most  related  species,  slightly  and  evenly  convex,  and  very  closely  and 
coarsely  granulated,  the  median  radial  and  submedian  rows  mostly 
hexagonal,  as  well  as  most  of  the  interradials.  Along  the  interradial 
margins  three  to  four  dorsal  plates  correspond  to  one  upper  mar- 
ginal. The  granules  are  coarse,  angular,  unequal  in  size,  flattish  or  a 
little  convex,  pretty  closely  in  contact,  mostly  eight  to  ten  on  the 
larger  plates. 

Small  valvular  pedicellarise,  commonly  a  little  larger  than  the 
granules,  but  often  of  about  the  same  size,  occur  on  a  few  of  the 
dorsal  plates,  on  the  marginals,  and  also  sparingly  on  the  actinals. 
The  valves,  which  in  the  larger  ones  project  a  little  above  the 
granules,  are  thin  and  flat,  about  as  high  as  broad.  They  are  mostly 
sunken  in  small,  roundish,  oblong  or  slightly  bilobed  pits. 

The  actinal  interradial  plates  .are  granulated  like  the  dorsals ;  they 
are  more  regular  in  form.  Adambulacral  plates  often  bear  a  bivalve 
pedicellaria;  they  have  about  six  short,  thick,  angular  spinules, 
crowded  in  two  rows  and  all  similar  in  height  and  size  to  the 
adjacent  granules.  The  inner  or  furrow  margin  bears  but  two,  or 
rarely  three,  short,  thick  spines. 

This  somewhat  resembles  C.  granularis,  but  its  granulation  is  much 
coarser,  its  margins  thicker  and  more  rounded,  and  its  distal  mar- 
ginal plates  more  irregular.  The  adambulacral  spines  are  much 


294  VERRILL 

coarser  and  fewer.  Madreporite  is  large.  Ambulacral  feet  hare 
large  terminal  suckers. 

It  was  known  to  me  only  from  the  Bering  Sea  and  Siberia.  The 
type  (U.  S.  Nat.  Mus.)  was  from  Bering  Island  (N.  Grebnitsky, 
1899). 

Fisher  (19116)  records  it  from  the  Commander  Islands,  Shumagm 
Islands,  Aleutian  Islands;  and  Kadiak,  Alaska,  from  low  tide  to 
1 02  fathoms. 

Genus  Amphiaster  Verrill. 

Amphiaster  VERRILL,  Trans.  Conn.  Acad.  Sci.,  i,  part  2,  p.  372,  February,  1878. 
Stellate,  with  short,  stout  rays  and  large  disk.  Dorsal  surface 
covered  with  large,  stout,  conical  spines  mostly  one  to  a  plate. 
Similar  spines  are  borne  by  most  of  the  marginal  plates  in  both 
series.  The  marginal  plates  are  unequal,  few,  large,  convex  or 
swollen,  and  naked  in  the  middle,  granulated  around  the  edges. 
Dorsal  abactinal  plates  are  granular,  polygonal,  tesselated,  sur- 
rounded by  papulae.  Interactinal  plates  tesselated;  each  usually 
bears  a  spine.  Adambulacral  plates  have  a  short  row  of  marginal 
spines  and  a  single  larger  spine  on  the  actinal  side.  Only  one 
species  is  known. 

AMPHIASTER  INSIGNIS  Verrill. 

Plate  xcvin,  figure  2  (type) . 

Amphiaster  insignis  VEBMLL,  Trans.  Conn.  Acad.  Sci.,  i,  part  2,  p.  372,  pi.  iv, 
fig.  10,  1878. 

A  well  grown  specimen  has  the  radii  50  mm.  and  24  mm. ;  ratio, 
about  i :  2.  The  marginal  plates  are  alternately  unequal ;  alternate 
ones  bear  a  large,  conical  spine.  There  are  about  ten  to  twelve 
lower  marginal  plates  and  eight  or  nine  upper  ones,  in  adult 
specimens. 

Three  pretty  regular,  radial  rows  of  spines  on  each  ray,  one  to 
each  of  the  principal  dorsal  plates ;  some  may  be  lacking.  Each  of 
the  interacting  plates  usually  bears  a  large  central  spine,  elsewhere 
they  are  coarsely  and  evenly  granulated. 

Lower  California;  La  Paz;  Gulf  of  California.  The  types  were 
from  La  Paz  (Yale  Museum). 

Subfamily  MEDIASTERINJS  Verrill,  1899. 
Stellate,  depressed,  in  form  resembling  Ceramaster.    Dorsal  radial 
plates  paxilliform,  or  parapaxillae,  with  circular  or  nearly  circular 
bases,  united  together  by  radiating  internal  ossicles,  about  six  to 


SHALLOW- WATER   STARFISHES  295 

each  plate,  producing  a  stellate  structure,  as  seen  from  the  inside. 
Supra-ambulacral  plates  are  present.  Pedicellariae  dorsal  and  ven- 
tral, sessile,  bivalve,  short  and  broad  in  the  type;  slender,  two-  or 
three-bladed  ones  may  occur  on  adambulacral  plates. 

Papulae  mostly  in  radial  rows,  emerging  from  between  the  con- 
nective radiating  ossicles,  often  two  or  three  together. 

The  adambulacral  spines  are  nearly  as  in  Ceramaster;  distal  ones 
are  larger  and  longer. 

The  principal  reason  for  distinguishing  this  subfamily  is  the  exist- 
ence of  radiating  internal  ossicles  connecting  the  bases  of  the 
adjacent  paxilliform  plates. 

This  seems  to  be  an  important  structural  feature,  and  in  the  case 
of  a  family  with  so  large  a  number  of  genera  as  Goniasteridae,  it  is 
advantageous  to  subdivide  it,  if  definite  and  important  structures  can 
be  found  for  subfamily  groups,  as  in  this  case. 

Genus  Mediaster  Stimpson. 

ifediaster  STIMPSON,  Journ.  Boston  Soc.  Nat  Hist,  n,  p.  490,  pi.  23,  figs. 

7-1 1,  1857.    Vert-ill,  Revision,  in  Trans.  Conn.  Acad.  Sci.,  x,  p.  178,  1899. 
Mediaster  SLADEN,  Voy.  Challenger,  Zool.,  xxx,  pp.  263,  752,  1889.     Fisher, 

op.  cit,  191  ib,  p.  196. 
Isaster  VERRILL,  Proc.  U.  S.  Nat  Mus.,  xvn,  p.  257,  1894. 

Form  stellate,  with  a  broad,  flat  disk  and  moderately  long,  tapered 
rays.  Marginal  plates  well  developed,  not  swollen,  granulated,  rather 
numerous,  higher  than  broad,  paired,  upper  and  lower  series  nearly 
equal  in  size  and  number,  and  with  their  sutures  more  or  less  corre- 
sponding vertically,  oblique  in  the  type.  No  odd  interradial  plate. 
Abactinal  plates  or  parapaxillae  are  regularly  longitudinally  arranged, 
of  moderate  size,  somewhat  elevated,  mostly  roundish,  covered  with  a 
rosette  of  short,  obtuse  spinules  or  elongated  granules.  When  these 
are  removed,  the  plates  on  the  central  part  of  the  disk  and  along  the 
median  region  of  the  arms  appear  as  roundish  or  oval  convex  bosses. 
They  are  connected  together  by  five  or  six  internal  radiating  ossicles, 
between  which  are  the  pores  for  the  papulae.  The  papulae  may  be 
single  or  (as  in  the  type)  clustered.  Thus  the  plates  appear  to  be 
stellate  at  the  base,  though  they  are  not  actually  of  that  shape.  The 
median  row  of  abactinal  plates  extends  to  the  apical  plate  of  the  rays 
in  the  type,  but  not  in  some  of  the  other  species.  Some  of  the  abac- 
tinal plates  bear  a  central  broad,  sessile,  valvular  pedicellaria,  which, 
in  the  type  species,  is  nearly  as  wide  as  the  plate.  Pedicellariae  are 
sometimes  lacking. 


296  VERRILL 

The  adambulacral  plates  bear  a  regular  marginal  row  of  three  to 
seven  slender  spinules,  and  usually  two  exterior  longitudinal  groups 
or  rows  of  shorter  spinules,  which  may  be  angular  and  obtuse ;  and 
toward  the  tips  of  the  rays,  some  of  them,  in  the  type,  become  larger 
and  longer,  as  in  Tosia  and  Ceramaster.  Some  of  these  spinules  may 
be  replaced  by  spinuliform  or  clavate,  two-  or  three-bladed  pedicel- 
lariae.  The  actinal  disk  plates  are  angular,  often  rhombic,  closely 
arranged  in  rows  parallel  with  the  ambulacral  grooves,  covered  with 
a  rosette  of  granules,  the  central  granules  often  replaced  by  a  wide 
valvular  pedicellaria.  The  dentary  plates  are  not  very  prominent; 
each  has  an  actinal  row  of  larger  spinules,  similar  to  those  of  the 
oral  margin. 

This  genus  closely  resembles  Ceramaster,  as  limited  above.  The 
principal  differences  consist  in  the  more  elevated  and  convex  abac- 
tinal  plates,  especially  in  the  papular  areas,  where  they  are  more 
widely  separated  by  the  large  papular  pores  and  united  by  interven- 
ing small  internal  ossicles,  which  give  them  a  stellate  appearance. 
On  other  parts  of  the  disk,  especially  near  the  interradial  margins, 
the  plates  are  angular  and  closely  joined  in  a  mosaic,  as  in  the  former 
genus.  The  large  valvular  pedicellariae  are  also,  to  some  extent,  char- 
acteristic, but  the  marginal,  actinal,  and  dentary  plates  and  their 
spinules  are  essentially  the  same  in  the  two  genera. 

MEDIASTER  ^EQUALIS  Stimpson. 
Plate  ii,  figure  i ;  plate  in,  figure  i ;  plate  v,  figures  3-5  (details). 

Mediaster  eequalis  STIMPSON,  Journ.  Boston  Soc.  Nat.  Hist.,  vi,  p.  490,  pi. 
xxiu,  figs.  7-1 1,  1857.  Whiteaves,  Trans.  Royal  Soc.  Canada,  iv,  p.  117, 
1887.  Verrill,  Trans.  Conn.  Acad.  Science,  i,  p.  326,  1867;  x,  p.  179,  pi. 
xxiv,  figs.  10-12,  1899.  Fisher,  op.  cit,  19116,  p.  198,  pi.  xxxv,  figs.  1-3; 
pi.  LIX,  figs,  i,  la-ic. 

Rays  five,  variable  in  length,  usually  about  equal  to  the  diameter  of 
the  disk,  regularly  tapered,  slender  at  the  tip.  Radii  usually  nearly 
as  1:3.  Marginal  plates,  on  each  side  of  a  ray  twenty-two,  above 
and  below,  in  a  specimen  having  the  greater  radius  36  mm.  The 
plates  on  the  margin  of  the  disk  are  higher  than  wide,  with  the 
intervening  sutures  somewhat  oblique.  The  lower  marginal  plates 
are  similar  in  size  and  shape.  All  are  closely  covered  with  small, 
rounded  granules.  Abactinal  areas  of  the  rays  are  wide  at  the  base, 
where  they  may  consist  of  seven  or  nine  rows  of  plates,  but  they 
rapidly  decrease  to  three  rows,  and  only  the  median  row  reaches  the 
apical  plate.  The  papular  areas  are  large,  covering  nearly  the  whole 


SHALLOW-WATER  STARFISHES  297 

width  of  the  proximal  half  of  the  rays,  as  well  as  most  of  the  central 
disk.  In  these  areas  the  plates  are  rounded  or  elliptical,  convex, 
somewhat  elevated,  and  separated  by  intervening  spaces,  in  which 
there  are  usually  five  or  six  groups  of  papular  pores,  the  individual 
pores  being  small  and  unequal,  two  or  three  or  more  forming  each 
group. 

Each  of  the  larger  abactinal  radial  plates  is  covered  with  a  rosette 
consisting  of  about  five  to  seven  central  and  twelve  to  fourteen  mar- 
ginal, short,  blunt,  or  clavate,  granule-like  spinules,  rather  longer 
than  broad.  Some  of  the  disk-plates  are  larger,  with  more  spinules. 
A  large  valvular  pedicellaria  often  replaces  the  central  group  of 
spinules  on  some  of  the  plates.  These  occupy  nearly  the  whole 
breadth  of  the  central  area  of  the  plate,  and  are  narrowly  oblong, 
not  much  elevated,  with  a  nearly  even  and  straight  margin.  Similar 
pedicellariae,  as  well  as  some  narrower  ones,  occupy  the  central  area 
of  some  of  the  interactinal  disk-plates. 

The  madreporic  plate  is  small,  sunken,  with  narrow,  acute  gyri. 
The  central  nephridial  pore  is  small  but  distinct. 

The  actinal  disk-plates  are  crowded  and  closely  united ;  those  next 
the  adambulacral  plates  are  squarish  or  rhombic  and  form  regular 
rows,  but  those  in  the  angles  are  smaller,  irregular,  and  more 
rounded.  All  are  covered  with  rosettes  of  granules,  or  short,  obtuse, 
often  prismatic  spinules,  rather  larger  and  less  regular  than  those  of 
the  upper  side.  A  central  valvular  pedicellaria  occurs  on  some  of 
the  plates,  as  stated  above. 

The  adambulacral  plates  are  squarish,  not  very  large.  Each  bears 
a  marginal  row  of  three  or  four  small,  oblong,  more  or  less  prismatic 
or  compressed,  blunt  spinules,  the  middle  one  usually  a  little  larger 
than  the  others.  External  to  these  are  two  sets  of  shorter  spinules, 
about  three  in  each  series ;  these  sometimes  form  two  rows,  but  in 
other  cases  are  in  a  rosette-like  group.  Those  next  the  inner  or 
groove-series  are  longer  than  the  others.  One  or  more  of  these, 
especially  distally,  may  be  replaced  by  a  spinuliform  pedicellaria 
with  two  or  three  blades.  On  the  distal  part  of  the  ray  one  or  two 
of  the  spinules  on  the  central  part  of  these  plates  becomes  consid- 
erably longer  and  larger  than  the  rest.  The  oral  spinules  are  similar 
to  the  adambulacral,  but  those  at  the  tip  of  the  oral  plates  are  rather 
larger  and  more  angular.  The  apical  plates  are  rather  small,  promi- 
nent, somewhat  obovate. 

The  preceding  description  is  from  a  Californian  specimen. 

I  have  examined  a  natural-size  photograph  of  the  type  of  this 
species,  furnished  by  Dr.  R.  Rathbun.  It  differs  from  the  specimen 


298  VERRILL 

described  above,  only  in  having  the  rays  somewhat  longer.  It  is 
poorly  preserved. 

This  species  is  closely  allied  to  M.  bairdii l  Verrill,  found  in  deep 
water  off  the  east  coast  of  the  United  States.  The  granulation  of 
both  surfaces  is,  however,  decidedly  finer  in  the  latter,  and  there  are 
various  other  differences  as  shown  by  the  figures. 

The  colors  in  life  are  very  bright.  The  dorsal  side  is  usually  deep 
red  or  vermillion ;  under  side  scarlet  or  orange,  varying  to  salmon- 
color;  ambulacral  feet  often  red  or  scarlet. 

Off  Wilmington,  California,  27  fathoms  (U.  S.  Nat.  Mus.,  speci- 
men described  above  and  figured)  ;  Queen  Charlotte  Islands,  Mal- 
colm Island,  low  water  (Whiteaves)  ;  Departure  Bay,  British  Colum- 
bia (C.  H.  Young,  Canadian  Geological  Survey,  1909)  ;  off  Victoria, 
Vancouver  Island  (C.  F.  Newcombe),  very  large. 

Fisher  records  it  from  numerous  localities,  in  6  to  167  fathoms, 
from  the  Alaskan  Peninsula  to  Lower  California.  Very  common  in 
20  to  50  fathoms. 

VARIATIONS. 

Two  specimens  from  Victoria,  British  Columbia,  1895,  received 
from  the  Provincial  Museum  through  C.  F.  Newcombe,  are  unusu- 
ally large,  and  have  an  appearance  quite  unlike  smaller  specimens 
usually  met  with. 

The  larger  has  the  radii  35  mm.  and  115  mm.;  dorsal  parapaxillae 
up  to  4  mm.  in  diameter.  The  other  has  the  radii  38  mm.  and 
103  mm.  The  rays  are  long  and  taper  to  slender  tips.  The  under 
surface  is  coarse  and  rough  in  appearance,  due  to  the  large  size  and 
the  length  of  the  adambulacral  spines  and  to  the  coarse  spinules  on 
the  interradial  plates. 

The  adambulacral  spines  are  about  the  same  in  number  and 
arrangement  as  on  the  smaller  specimens,  but  are  much  increased  in 
relative  size ;  they  are  up  to  3.5  mm.  long.  The  same  is  true  of  the 
interradial  spinules.  These  last  stand  in  stellate  groups,  mostly  of 

1 MEDIASTER  BAIRDII  Verrill.    (Plate  n,  figure  2;  plate  in,  figure  a.) 
Archaster  bairdii  VERRILL,  Amer.  Journ.  Sci.,  xxm,  p.  139,  1882. 
Isaster  bairdii  VERRILL,  Proc.  U.  S.  Nat  Mus.,  xvn,  p.  258,  1894.     Amer. 

Journ.  Sci.,  XLIX,  p.  136,  1895. 
Mediaster  stellatus  PERRIER,  Mem.  Soc.  de  France,  IT,  p.  268,  1891.    Resultats 

des  Campag.  Scient.,  fas.  xi,  p.  46,  1896,  pi.  IT,  figs,  i-irf. 
Mediaster  bairdii  VERRILL,  Trans.  Conn.  Acad.,  x,  p.  181,  pi.  xxnr,  figs.  1-9, 

pi.  xxv,  figs.  8,  8a,  1898. 
Off  the  east  coast  of  the  United  States,  in  351  to  721  fathoms. 


SHALLOW-WATER   STARFISHES  299 

seven  to  nine,  but  often  in  a  circle  with  one  or  two  central  on  each 
plate.  They  are  short,  thick,  mostly  angular,  often  three-sided, 
mostly  blunt,  but  many  are  acute  and  often  beech-nut  shaped.  The 
central  one  is  frequently  replaced  by  a  short,  thick,  bivalved,  or 
sometimes  trivalved,  erect  pedicellaria.  In  the  larger  specimen  the 
spinules  are  erect  in  a  close,  round  cluster,  leaving  open  intervals 
between.  In  the  other  they  are  divergent,  covering  most  of  the 
interspaces.  The  inner  adambulacral  spines  form  a  nearly  straight, 
close  row  of  three  or  four,  which  are  nearly  equal  in  length  and 
more  or  less  angular  and  blunt ;  the  middle  row  usually  consists  of 
three  or  four  similar  but  shorter  spines  in  a  curved  or  oblique  series ; 
outside  of  these,  near  the  outer  margin  of  the  plate,  are  four  to  six 
much  shorter  angular  spinules  in  a  curved  group,  like  those  on  the 
interradial  plates.  A  bivalve  pedicellaria  sometimes  replaces  some 
of  these.  In  a  few  cases  all  the  spinules,  except  those  of  the  inner 
row,  form  a  circular,  convergent  group  of  about  six  nearly  equal 
spines. 

The  dorsal  parapaxillae  are  very  regular,  circular,  or  broad-ellipti- 
cal, evenly  covered  with  large,  rounded  granules.  About  twenty 
to  twenty-five  granules  form  an  outer  circle  on  the  larger  plates; 
inside  these  is  a  second  circle,  usually  of  eight  to  twelve  somewhat 
larger  granules  on  the  larger  plates,  or  about  six  on  the  smaller 
ones;  the  center  is  occupied  by  a  single  round  granule  of  similar 
size,  or  by  two  or  three  on  the  larger  plates.  Sometimes  a  small, 
stout,  erect  bivalve  pedicellaria  occupies  the  center.  Sometimes  it 
is  one-third  as  broad  as  the  plate. 

The  marginal  plates  are  evenly  covered  with  close,  rounded  gran- 
ules; the  marginal  ones  are  a  little  larger  and  divergent  over  the 
margins,  thus  forming  narrow  covered  channels  between  the  plates. 
Some  of  the  marginals  bear  small  bivalve  pedicellariae,  like  those  of 
the  dorsal  plates. 

In  one  of  these  specimens  the  rays  are  considerably  bent  laterally, 
and  some  of  them  are  twisted  and  have  the  tips  upturned,  showing 
more  flexibility  than  the  heavy  plating  would  be  expected  to  permit 

Subfamily  HIPPASTERIINJE  Verrill. 
Hippasteriina  VERRILL,  Revision  Genera,  etc.,  in  Trans.  Conn.  Acad.,  x,  p. 

174,  1899.    Fisher,  op.  cit,  19116,  p.  223. 
Antheneida  (pars)  PERRIER  and  others. 

This  group  includes  stout,  pentagonal  or  short-rayed  starfishes, 
that  have  large,  elongated,  divergent,  and  differentiated  adambula- 
cral spines,  one  or  two  larger  ones  situated  on  the  central  part  of 


30O  VERRILL 

each  plate.  The  dorsal  and  marginal  plates  are  large,  convex,  bor- 
dered or  covered  with  large  granules,  and  often  have  one  or  more 
central  tubercles  or  stout  spines ;  no  fascioles. 

Bivalve  pedicellariae  having  short,  broad  valves,  and  often  of  large 
size,  are  usually  present,  both  on  the  dorsal  and  actinal  plates,  and 
often  on  the  marginals ;  smaller  spatulate  forms  also  occur. 

The  abactinal  plates  are  thick,  rather  closely  joined,  and  polygonal 
or  roundish,  usually  unequal,  with  small  intermediate  secondary 
plates. 

The  actinal  plates  are  numerous,  tesselated,  and  covered  with 
large  granules,  at  least  around  the  margin,  and  usually  have  either 
a  large  valvular  pedicellaria  or  else  a  short,  thick  spine  or  tubercle 
in  the  middle.  Superambulacral  plates  are  lacking. 

In  this  subfamily  I  include  Cladaster  Verrill,  not  included  by 
Fisher.  It  lacks  the  large  dorsal  and  marginal  spines  or  conical 
tubercles  of  Hippasteria,  and  the  dorsal  ossicles  are  all  of  one  kind. 
Fisher  describes  (ignb,  p.  22,  pi.  XLI,  figs,  i,  2)  C.  validus,  Fisher, 
from  off  the  Aleutian  Islands,  in  283  fathoms. 

Genus  Hippasteria  Gray. 

Plate  XLVII  ;  plate  XLVIII,  figures  1-5  (details  of  type  species)  ;  plate  XLIX, 

figure  6. 

Goniaster  (pars)  AGASSIZ/  Prodromus,  Mem.  Soc.  Sci.  Neuchatel,  i,  p.  191, 
1835  (non  Gray,  1840).  Forbes,  Brit.  Starfishes,  p.  122,  1842.  Norman, 
Ann.  and  Mag.  Nat.  Hist.,  xv,  p.  123,  1865. 

Hippasteria  GRAY,  Ann.  and  Mag.  Nat.  Hist.,  vi,  p.  279,  1840;  Synopsis,  p.  9, 
1866. 

Hippasteria  PERKIER,  Arch.  Zool.  Exper.,  v,  p.  86,  1876.  Viguier,  Arch.  Zool. 
Exper.,  VH,  p.  176,  1878  (structure).  Sladen,  Voy.  Challenger,  Zool.,  xxx, 
p.  341,  1889.  Verrill,  op.  cit,  1899,  p.  174,  Fisher,  op.  cit.,  191 1&,  p.  223. 

Astrogonium  (pars)  MULLER  and  TROSCHEL,  System  Asteriden,  p.  52,  1842. 

Hippasterias  BELL,  Catal.  Brit.  Echinod.,  p.  76,  1892  (spelling  incorrect). 

Disk  broad,  rather  flat,  subpentagonal,  with  short  rays,  and  con- 
cave interradial  margins.  Marginal  plates  of  moderate  size,  thick, 
convex,  regularly  paired,  surrounded  by  a  marginal  row  of  granules 
and  usually  bearing  one  or  more  short,  stout  spines  or  tubercles,  and 
sometimes,  also,  a  wide,  bivalvular  pedicellaria.  Dorsal  or  abactinal 
plates  mostly  large,  more  or  less  convex,  thick,  tesselate,  subpolyg- 
onal  or  roundish,  with  marginal  granules,  and  bearing  one  or  more 
thick  central  spines  or  tubercles,  or  else  a  wide  bivalved  pedicellaria ; 
many  small  secondary,  granulated  plates  are  interspersed  between 

1  See  note,  p.  286,  for  original  contents  of  Goniaster  Agassiz. 


SHALLOW-WATER   STARFISHES  3OI 

the  larger  ones,  and  may  not  have  the  central  spine.  Plates  some- 
times partially  obscured  by  a  thin  integument. 

Actinal  plates  rather  large,  angular,  closely  tesselated,  arranged  in 
rows  parallel  with  the  adambulacrals.  They  are  surrounded  by  a 
row  of  coarse  granules  and  bear  either  a  large  and  wide  bivalved 
pedicellaria,  like  those  of  the  dorsal  plates,  or  else  one  or  more 
central  tubercles.  Sometimes  nearly  all  of  them  bear  large  pedi- 
cellariae. 

Adambulacral  plates  large,  but  smaller  than  the  interactinals, 
squarish,  bearing  a  marginal  row  of  large  granules,  or  short,  blunt 
spinules,  one,  or  sometimes  two,  stout,  central  spines,  and  a  longi- 
tudinal furrow-series  of  few,  usually  two  or  three,  smaller  divergent 
spines.  (See  pi.  XLVIII,  fig.  5.) 

Oral  or  dentary  plates  large,  bearing  a  marginal  row  of  rather 
strong  oral  spines,  besides  two  rows  of  shorter  epioral  ones  on  the 
actinal  surface.  (See  pi.  XLVIII,  fig.  2.) 

Median  dorsal  pore  large,  surrounded  by  papillae.  ( See  pi.  XLVIII, 
fig.  4.)  Tube- feet  stout,  with  large  suckers. 

Madreporic  plate  rather  small,  with  few  coarse  gyri.  (PL  XLJX, 
fig.  6.)  Papulae  small,  isolated,  in  the  angles  between  the  abactinal 
plates.  (See  pi.  XLVIII,  figs.  1-4.) 

Besides  the  following  species,  Professor  Fisher  has  described  three 
deep-water  species  from  the  North  Pacific.  Neither  of  them  has 
been  recorded  from  within  the  limits  of  this  report. 

The  North  Atlantic  and  Arctic  species,  H.  phrygiana,  common  on 
the  coast  of  New  England,  in  50  to  200  fathoms,  has  not  yet  been 
found  on  the  Pacific  side.  (See  pi.  XLVII;  pi.  XLVIII,  figs.  1-5;  pi. 
XLIX,  fig.  6,  for  details.) 

HIPPASTERIA  SPINOSA  Verrill. 
Plate  L,  figures  4-4^;  plate  xcvm,  figure  i  (type). 

Hippasteria  spinosa  VERRILL,  Amer.  Journ.  Sci.,  xxvm,  p.  63,  July,  1909. 
Fisher,  op.  cit,  191  ib,  p.  224,  pi.  XLII,  figs.  1-3;  pi.  XLIII,  figs,  i,  2;  pi.  uc, 
fig-  4- 

Arms  rather  longer  than  in  the  Atlantic  species.  Upper  surface 
thickly  covered  with  rather  slender,  tapered,  acute,  unequal  spines, 
nearly  every  plate  bearing  a  single  spine ;  those  that  do  not  have  a 
spine  carry  a  large  central  pedicellaria,  about  equal  in  diameter  to  the 
corresponding  spine,  but  varying  in  size,  like  the  spines,  according 
to  the  size  of  the  plate.  The  spines  and  pedicellariae  are  surrounded 
at  the  base  by  one  or  more  circles  of  granules  borne  on  the  plates 


3O2  VERRILL 

around  the  margins,  leaving  the  mammilliform  central  part  naked. 
The  dorsal  pedicellariae  are  oblong-elliptical,  as  seen  from  above ;  the 
jaws  are  convex,  usually  a  little  wider  distally  than  at  the  base, 
often  nearly  as  high  as  wide.  The  upper  marginal  plates  mostly 
bear  two  divergent  spines,  similar  to  those  of  the  back.  The  lower 
marginal  plates  also  bear  similar,  but  shorter,  spines,  usually  two, 
but  sometimes  one  or  three. 

The  actinal  plates  are  coarsely  granulated  and  each  usually  bears 
a  large  central  pedicellaria  similar  to  those  of  the  dorsal  surface  in 
size  and  form,  but  usually  rather  higher  and  with  the  edges  of  the 
jaws  distinctly  denticulated. 

Color,  as  dried,  orange  above,  yellow  below. 

Greater  radius,  80  mm.,  lesser  radius,  40  mm. 

The  type  specimen  was  taken  at  Departure  Bay,  British  Colum- 
bia, in  18  fathoms,  gravel,  September,  1908  (Mr.  C.  H.  Young, 
Canadian  Geological  Survey). 

A  smaller  specimen  from  Puget  Sound,  sent  by  Professor  Kincaid, 
has  the  radii  28  mm.  and  65  mm.  It  agrees  very  well  with  the 
larger  one  in  the  number  and  form  of  the  dorsal  pedicellariae  and 
spines,  but  the  latter  are  slightly  less  acute.  The  marginal  plates 
of  both  series  mostly  bear  two  spines,  like  the  dorsals,  but  rather 
larger,  while  there  are  often  two  or  three  additional,  smaller,  obtuse 
spines  on  the  lower  ones.  The  pedicellariae  of  the  actinal  plates  are 
numerous,  about  as  high  as  long,  and  similar  in  size  to  the  dorsals. 
The  adambulacral  spines  are  mostly  not  preserved.  At  the  distal 
third  there  are  two  furrow-spines  to  a  plate. 

The  specimens  of  this  species  described  by  me  are  from  Puget 
Sound  (Kincaid)  and  Departure  Bay,  British  Columbia  (Canadian 
Geological  Survey). 

Mr.  Fisher  (19116)  records  it  from  many  localities,  from  Kadiak, 
Alaska,  to  Southern  California,  in  27  to  121  fathoms. 

Family  ODONTASTERID&  Verrill. 

Odontasteridce  VERRILL,  Trans.  Conn.  Acad.  Sci.,  x,  p.  201,  1899.    Fisher,  1905, 

p.  302;  191 1  b,  p.  153. 
Gnathasterina  (Pars)  PERKIER,  1894,  p.  251. 

Form  stellate  with  broad,  short  rays,  or  pentagonal.  Marginal 
plates  prominent,  with  an  odd  interradial  plate  in  each  row,  nor- 
mally. 

The  jaws  bear  one  or  two  erect  or  recurved  hyaline  spines  on  the 
actinal  (epioral)  side,  near  the  apex. 


SHALLOW-WATER   STARFISHES  303 

Abactinal  plates  are  more  or  less  paxilliform,  parapaxillae  or  pro- 
topaxillae,  with  intervening  large  papular  pores  on  the  radial  areas. 
The  abactinal  plates  may  bear  clusters  of  more  or  less  elongated 
spines,  or  a  group  of  small  granules.  They  usually  form  obliquely 
transverse  lines  on  the  rays,  not  always  regular. 

Interactinal  plates  angular,  covered  either  with  spines  or  with 
granules.  Small,  simple  pedicellariae  sometimes  occur  on  the  actinal 
or  abactinal  plates.  They  may  have  two,  three,  or  four  papilliform 
blades. 

Adambulacral  plates  usually  bear  elongated  spinules,  arranged 
in  three  or  four  small  transverse  rows ;  generally  only  two  or  three 
in  the  furrow-series;  sometimes  only  one.  Dentary  plates  usually 
have  elongated,  acute  marginal  and  apical  spines.  They  are  some- 
times closely  united  along  the  median  suture ;  in  other  cases  ( Odont- 
cater)  they  are  separated  by  a  space  covered  only  by  membrane. 

The  marginal  plates  are  covered  either  with  spinules  or  with 
granules,  sometimes  the  upper  ones  are  granulated  and  the  lower 
spinulose,  like  the  corresponding  disk-plates ;  they  usually  have  deep- 
fasciolatcd  sutures. 

Ambulacral  feet  have  suckers.    No  superambulacral  plates. 

Genus  Odontaster  Verrill. 

Odontaster  VERJULL,  Amer.  Journ.  Sci.,  xx,  p.  402,  1880 ;  Proc.  U.  S.  Nat.  Mus., 
xvn,  p.  262,  1894;  Amer.  Journ.  Sci.,  XLIX,  p.  136,  1897;  op.  cit,  1899, 
p.  205. 

Gnathaster  PERKIER  (pars),  Exp.  Trav.  et  Talism.,  p.  244,  1894. 

Odontaster  BELL  (pars),  Proc.  Zool.  Soc.  London,  p.  260,  1893.  Fisher, 
op.  cit.,  i8n&,  p.  154. 

A  single,  odd,  hyaline,  recurved,  movable  spine  on  the  apex  of 
each  jaw.  Dentary  plates  large,  separated  by  an  open,  fusiform 
space,  covered  by  membrane.  Abactinal  surface  covered  with  ele- 
vated, convex,  or  clavate  paxilliform  plates,  or  parapaxillae,  which 
usually  bear  clusters  of  elongated  spinules,  like  true  paxillae;  their 
bases  are  stellate.  Upper  marginal  plates  are  usually  finely  spinu- 
lated.  Lower  marginal  plates  are  generally  large  and  placed  singly 
or  in  the  angles  around  the  radial  paxillae.  The  radial  abactinal 
plates  form  more  or  less  evident  obliquely  transverse  rows  and 
extend  nearly  or  quite  to  the  apical  plate. 

The  odd  interradial  marginal  plate  is  usually  triangular  or  wedge- 
shaped.  Simple  pedicellariae  occur  rarely. 

The  adambulacral  plates  usually  bear  several  rows  of  spines; 
usually  three  or  four  in  the  furrow-series,  rarely  but  two. 


304  VERRILL 

The  open  suture  between  the  dentary  plates  of  the  jaws;  the 
movable  hyaline  spine,  attached  only  by  its  base,  at  the  apex  of  the 
jaw,  together  with  the  very  spinose  character  of  the  abactinal 
paxillae  and  marginal  plates,  separate  this  genus  from  its  allies.  The 
marginal  plates  are  also  larger  than  in  most  of  the  other  groups, 
and  the  adambulacral  plates  bear  usually  three  or  four  spines  in 
the  furrow-series. 


ODONTASTER  CRASSUS  Fisher. 

Odontaster  crassus  FISHER,  op.  cit,  1905,  p.  302;  op.  cit.,  191  ib,  p.  154,  pi. 
xxix,  figs.  1-4;  pi.  LW,  fig.  6. 

Form  stellate  with  short  rays;  radii  of  the  type,  13  mm.  and 
21  mm.;  ratios,  about  1:1.5.  Marginal  plates  few,  massive,  eight 
in  a  row  in  the  type,  covered  with  granule-like  spinules.  Odd 
epioral  hyaline  spine  is  lanceolate,  recurved.  Interactinal  plates 
squarish,  bearing  five  to  twelve  radiating,  short,  stout,  pointed 
spinules. 

Dr.  Fisher  records  it  from  ten  stations,  in  43  to  284  fathoms,  from 
off  Monterey  to  San  Diego,  California. 

Family  ASTEROPIDM  Fisher. 

Gymnasteriida  PERKIER,  1884,  pp.  165,  229.  Sladen,  1889,  p.  355. 
Gytnnasteriidce  -\-  Poraniidce  PERRIER,  op.  cit.,  1894,  pp.  163,  227.* 
Asteropidte  FISHER,  1908,  p.  90;  19116,  p.  247. 

Disk  usually  large;  rays  short  and  broad.  Dorsal  ossicles  some- 
times tesselated,  in  regular  radial  rows,  more  often  irregular,  some- 
times partly  abortive,  sometimes  reticulated;  either  covered  with  a 
thick,  smooth  or  granulated  skin,  or  spinose,  or  spinulose. 

Marginal  plates  various,  sometimes  prominent,  usually  oblique  or 
overlapping,  sometimes  with  a  small  group  of  spinules  on  the  outer 
edge,  or  with  a  single  spine;  sometimes  nearly  abortive,  and  the 
edge  of  disk  thin. 

Interactinal  plates  sometimes  numerous,  usually  in  regular  oblique 
rows.  Papulae  in  dorsal  radial  areas,  sometimes  intermarginal. 
Pedicellariae  commonly  lacking;  when  present,  valvular,  with  two  to 
four  valves. 

*It  seems  to  the  writer  very  doubtful  whether  the  Poraniidae  should  be 
combined  with  this  family.  Some  of  the  genera  are  very  similar  to  Asterinidae 
in  form  and  structure. 


SHALLOW-WATER   STARFISHES  305 

Subfamily  ASTEROPIN1E  Verrill,  nov. 

Margins  not  thin ;  upper  and  lower  marginal  plates  well  developed, 
subequal.  Dorsal  skeleton  well  developed,  the  plates  tesselated  or 
stellate-reticulate,  with  or  without  spinules  or  spines.  Superomar- 
ginal  plates  sometimes  with  stout  spines.  Interradial  plates  numer- 
ous, in  chevrons.  Adambulacral  spines  few.  Pedicellariae,  when 
present,  bivalve  or  multivalve. 

The  Poraniinae  differ  in  having  the  marginal  plates  feebly  devel- 
oped, more  unequal,  the  edges  of  the  disk  being  thin  in  most  cases, 
and  formed  by  the  inf eromarginals ;  the  dorsal  plates  are  irregular, 
often  partly  or  nearly  all  abortive,  not  forming  a  regular  median 
row,  and  usually  concealed  by  a  thick,  smooth,  or  else  finely  spinu- 
lose  dermis;  interactinal  plates  fewer,  in  oblique  rows,  or  nearly 
abortive,  or  in  detached,  transverse  rows  (  Tylaster) ,  with  or  without 
spinules.  Pedicellariae  generally  lacking. 

Genus  Dermasterias  Perrier. 

Dermasterias  PERKIER,  Revision  Stell.,  Arch,  de  Zool.  Exper.,  v,  p.  98,  1876. 

Viguier,  op.  cit,  vn,  p.  218,  1878.     Sladen,  Voy.  Chall.,  Zool.,  xxx,  pp. 

355.  375,  1889.    Fisher,  op.  cit.,  19116,  248. 
Asteropsis  (pars')  GRUBE,  Wieg.  Arch.,  xxm,  1857.    A.  Agassiz,  North  Amer. 

Starfishes,  Mem.  Mus.  Cotnp.  Zool.,  v,  p.  106,  pi.  xv  (structure). 

Some  of  the  characters  of  this  genus  have  not  been  correctly 
stated  by  several  previous  writers. 

It  has  more  or  less  numerous  small  pedicellariae  scattered  irregu- 
larly over  the  dorsal  surface.*  These  are  partly  bivalve  and  partly 
trivalve,  while  four-valved  ones  often  occur.  The  valves  are  small, 
thick,  obtuse  or  rounded.  Five-valved  ones  are  rarely  seen ;  three- 
valved  ones  are  often  the  most  numerous.  Other  pedicellariae,  of 
much  larger  size,  often  occur  just  outside  of  the  bases  of  the  inner 
adambulacral  spines.  They  may  stand  in  a  single  row,  or  they  may 
be  crowded  into  two  or  three  rows.  (PI.  vi,  figs.  4,  5.)  They  are 
mostly  bivalvular,  with  wide,  short,  truncate  jaws,  but  some  of  them 
have  three  jaws  (pi.  vi,  fig.  5,  Pf  P7)  ;  occasionally  one  occurs  with 
four  smaller  jaws. 

On  one  of  the  young  specimens,  they  nearly  cover  the  surface, 
and  most  of  them  have  four  valves;  some  have  three,  and  a  few 
five  valves.  (See  pi.  L,  figs,  i-ifr.) 

1  Perrier  (ReVis.  Stell.,  p.  98,  1876)  gives  the  absence  of  pedicellariae  as  a 
character  of  the  genus.     On  some  specimens  from  Puget  Sound  they  are  very 
numerous  on  the  dorsal  side  (var.  valvulifera  Verrill). 
21 


306  VERRILL 

The  adambulacral  spines,  which  are  usually  two  to  a  plate,  inner 
and  outer,  increase  in  length  and  become  terete,  instead  of  flat,  at 
the  tips  of  the  arms,  while  some  of  the  distal  interactinal  and  inferior 
marginal  plates  sometimes  bear  short,  thick,  stumpy  spines,  especially 
in  large  specimens. 

In  formalin  preparations  the  entire  surface  is  covered  with  a 
thick,  soft  skin,  deep  brown  or  yellowish  brown  in  color,  which 
entirely  conceals  the  ossicles  and  spines  (pi.  xxix,  fig.  2),  but  the 
groups  of  papulae  on  the  back  are  quite  distinct.  In  life  this  skin  is 
very  soft  and  covered  with  abundant  mucus.  In  some  dried  prepa- 
rations the  integument  is  like  a  sort  of  thin  varnish  over  the  surface, 
leaving  the  ossicles  easily  visible,  but  in  other  cases  it  is  so  dried 
that  the  ossicles  are  mostly  concealed.  The  principal  dorsal  ossicles, 
both  on  the  rays  and  on  the  disk,  have  a  regular  stellate  arrangement, 
as  stated  by  Agassiz.  Each  large  ossicle  is  roundish,  with  slight 
lobes,  connected  with  five  to  eight  small,  radiating  connective 
ossicles.  In  the  interradial  regions  the  ossicles  are  stouter,  are  not 
stellate,  and  are  closely  imbricated.  In  this  area  there  are  few  or  no 
papulae.  The  interactinal  plates  are  broad  ovate,  and  form  rows 
parallel  to  the  ambulacra.  They  are  distinctly  alternatingly  imbri- 
cated, like  slates  on  a  roof.  Papulae  are  very  numerous  and  form 
large  groups  on  the  dorsal  side  of  large  specimens,  but  stand  singly 
on  young  ones. 

There  is  an  incomplete,  partly  calcified  interradial  septum,  extend- 
ing outward  from  the  jaws,  composed  of  rather  large  rounded 
ossicles.  Superambulacral  plates  ( ?)  of  small  size  are  present.  For 
some  details  of  the  skeleton  see  pi.  LXXXVI,  figs.  2-2d. 

Only  one  species  of  the  genus  is  known. 

DERMASTERIAS  IMBRICATA  (Grube)  Perrier. 

Plate  vi,  figures  3,  4,  5;  plate  xxix,  figure  2;  plate  L,  figures  i-ib  (pedicel- 
lariae)  ;  plate  LXXXVI,  figures  2-2d  (skeleton)  ;  plate  xcvii,  figures  2-2b 
(variety). 

Asteropsis  imbricata  GRUBE,  Wieg.  Archiv,  xxm,  p.  340,  1857.  Verrill,  Trans. 
Conn.  Acad.  Sci.,  i,  p.  324,  1867  (distribution).  A.  Agassiz,  North  Ameri- 
can Starfishes,  p.  106,  pi.  xv,  figs.  1-7,  1877  (structure). 

Dermasterias  inermis  PERKIER,  Revision  Stellerides,  p.  282,  1875. 

Dermasterias  imbricata  PERRIER,  Arch,  de  Zool.  Exper.,  v,  p.  98,  1876.  Viguier, 
op.  cit.,  vn,  p.  218,  1878.  Sladen  Voy.  Chall.,  xxx,  p.  766,  1889.  Whiteaves, 
Trans.  Royal  Soc.  Canada,  iv,  p.  117,  1887.  Clark,  Proc.  Boston  Soc. 
Nat.  Hist,  xxix,  p.  325,  pi.  i,  figs,  i,  2,  1901.  Fisher,  op.  cit,  191  ib, 
p.  249,  pi.  XL,  figs,  i,  2 ;  pi.  LVI,  fig.  i. 


SHALLOW-WATER   STARFISHES  307 

The  disk  is  large,  rather  plump  in  life,  with  five  short,  tapering 
rays  (rarely  six),  but  the  length  of  the  rays  is  subject  to  considerable 
variation.  The  radii  of  a  short-rayed  Alaskan  specimen  measure 
50  mm.  and  120  mm.  A  long-rayed  one,  from  the  same  place, 
measures  35  mm.  and  no  mm. 

Surface  smooth  and  lubricous  in  life,  owing  to  the  thick,  soft 
skin.  When  dried,  the  ossicles  are  more  or  less  distinctly  visible 
on  both  sides.  Those  of  the  back  have  a  rather  regular  stellate 
arrangement  on  the  disk  and  bases  of  the  rays. 

Many  specimens  have  numerous  small,  scattered,  dermal  dorsal 
pedicellariae,  mostly  bivalve  or  trivalve,  but  often  quadrivalve,  as 
described  above,  under  the  genus.  In  many  specimens  they  are 
lacking. 

Larger  ventral  pedicellarise  are  often  present.  In  most  cases  they 
form  a  single  row,  more  or  less  concealed  under  the  free  edge  of  the 
dermal  membrane  in  dried  specimens.  In  nearly  all  cases  they  take 
the  place  of  the  outer  row  of  adambulacral  spines,  for  they  occupy 
the  same  position  on  the  plate.  Spines  are  absent  when  pedicellariae 
are  present ;  sometimes  the  latter  will  form  regular  rows  for  half  the 
length  of  the  grooves,  and  then  spines  will  take  their  place  on  the 
distal  half,  or  they  may  alternate  irregularly  with  spines.  The 
breadth  of  these  pedicellariae  is  almost  the  same  as  that  of  the  spines, 
but  they  are  not  half  as  long.  Many  specimens  lack  these  ventral 
pedicellariae  entirely. 

In  one  lot  of  twenty-three  dried  specimens  from  Sitka,  four  had 
few  or  no  dorsal  pedicellariae ;  nineteen  had  dorsal  pedicellariae ;  five 
of  these  had  large  numbers ;  twelve  had  them  both  on  the  dorsal  and 
on  the  ventral  side ;  five  had  numerous  actinal  ones. 

Their  presence  does  not  depend  upon  age,  for  they  are  lacking 
on  some  of  the  largest  specimens,  while  they  are  numerous  on  others. 
They  are  abundant  on  some  of  the  young  ones,  about  60  mm.  in 
diameter. 

The  color,  in  life,  according  to  Prof.  W.  R.  Coe,  is  deep  orange 
to  dull  red. 

One  small,  six-rayed  example  was  found  by  Professor  Coe  (pi.  vi, 
fig-  3)  ;  its  ra-dii  are  7  mm.  and  14  mm. 

This  species  is  common  from  Monterey  Bay,  California,  to  middle 
Alaska.  Numerous  specimens  were  collected  at  Sitka,  etc.,  by  Prof. 
W.  R.  Coe,  on  the  Harriman  Expedition.  It  lives  at  and  just  below 
low  tide.  It  is  also  common  in  Puget  Sound,  whence  I  have  seen 

»many  specimens.    Several  large  specimens  have  been  sent  to  me  by 


308  VERRILL 

the  Canadian  Geological  Survey,  from  British  Columbia  and  the 
Queen  Charlotte  Islands,  Vancouver  Island,  etc.  It  has  not  been 
recorded  except  from  very  shallow  water  or  between  tides.  Dr. 
Fisher's  records  are  from  Sitka  to  Monterey,  California,  shores, 

DERMASTERIAS  IMBRICATA,  var.  VALVULIFERA 
Verrill,  nov. 

Plate  xcvii,  figures  2-26. 

This  name  is  proposed  for  the  form  of  this  species  having  many 
(often  very  numerous)  valvular  pedicellariae  on  the  dorsal  surface. 
Usually  there  is  also  a  row  on  the  actinal  side,  external  to  and 
parallel  with  the  inner  adambulacral  spines,  while  others  may  occur 
on  the  interactinal  areas,  either  centrally  or  more  often  proximally, 
near  the  jaws,  in  pairs.  The  actinal  pedicellariee  mostly  have  two 
broadly  rounded  jaws,  sometimes  three ;  those  of  the  dorsal  surface 
mostly  have  three  or  four  narrower  valves,  but  some  may  have  two ; 
others  five  or  six  jaws  (see  pi.  xcv,  figs.  2-2&).  The  specimen 
figured  from  Sitka  has  the  radii  24  mm.  and  38  mm. ;  another  from 
the  same  place  has  the  radii  38  mm.  and  77  mm.  Larger  specimens 
occur. 

Except  for  the  presence  of  many  pedicellariae  this  variety  does  not 
differ  much  from  the  ordinary  variety  with  few  or  no  pedicellariae. 
This  does  not  depend  upon  age,  for  both  large  and  small  specimens 
occur  of  each  form.  Very  young  specimens,  as  usual  in  other  genera, 
lack  pedicellariae. 

All  the  specimens  seen  are  from  southern  Alaska,  at  Sitka,  etc. 

Family  OPHWIASTERW2E  Verrill. 

Ophidiasteridee  VERRILL,  Trans.  Conn.  Acad.  Sci.,  I,  part  2,  p.  344,  1867. 
Linckida  PERRIER,  Revision  Stellerides,  p.  117,  1875. 

Linckiida  SLADEN  (pars),  op.  cit.,  1889,  p.  397.  Fisher,  op.  cit.,  191  ib,  p.  240. 
Linckiada  VIGUIER,  Squelette  des  Stell.,  p.  144,  1879. 

Form  stellate,  usually  with  five  slender  rays,  but  the  number  is 
variable  in  some  autotomous  species.  Marginal  plates  usually  small 
and  not  conspicuous.  Abactinal  and  marginal  ossicles  usually  tes- 
selated  or  subimbricated,  in  most  cases  granulose  (covered  with 
rather  thick,  smooth  skin  in  Leiaster},  rarely  spinose. 

Papulae  usually  numerous,  mostly  abactinal  and  lateral  in  groups, 
sometimes  single;  in  the  majority  of  the  genera  they  occur  also 
below  the  marginals,  or  between  thp  interactinal  plates. 

Adambulacral  plates  with  either  granules  or  spinules. 


SHALLOW- WATER   STARFISHES  309 

Pedicellariae  often  lacking;  when  present,  usually  bivalve,  some- 
times spatulate  and  excavate  or  fossate.  Superambulacral  plates 
often  present. 

Genus  Linckia  Nardo. 

Linckia  NARDO,  Oken's  Isis,  1834,  p.  717.  Gray,  op.  cit.,  1840,  p.  284;  System, 
p.  13,  1866.  Verrill,  op.  cit.,  1867,  p.  285.  Sladen,  op.  cit,  1859.  Fisher, 
op.  cit,  191  ib,  p.  242. 

Ophidiaster  (pars)  MULLER  and  TROSCHEL,  Syst,  p.  28,  1842. 

Phataria  MONKS,  op.  cit.,  1903,  p.  35  (non  Gray). 

Disk  small,  rays  long,  slender,  nearly  terete,  usually  variable  in 
number.  Some  species  are  autotomous.  Adambulacral  plates  bear 
granule-like  structures,  in  two  series.  Papulae  lacking  on  actinal  side, 
numerous  in  abactinal  areas.  Abactinal  plates  irregularly  arranged. 
Pedicellariae  not  observed. 

LINCKIA  COLUMBLE  Gray. 

Linckia  Columbia  GRAY,  op.  cit.,  1840,  p.  285;  Synopsis,  1866,  p.  14.     Sladen, 

op.  cit.,  1889,  p.  784.    Verrill,  op.  cit,  1867,  pp.  332-334.    Fisher,  op.  cit, 

ignb,  p.  242,  pi.  XLVIII,  figs.  1-7. 
Linckia  diplax  PERRIER,  Revision  Stell.,  p.  144,  1875. 
Phataria  (Linckia)  fascialis  MONKS,  op.  cit,  1003,  LV,  p.  351  (autotomy  and 

variations). 
Phataria  (Linckia)  unifascialis,  var.  bifascialis  MONKS  (non  Gray)  ;  op.  cit., 

1904,  LVI,  p.  596,  pi.  XLII  (autotomy,  variations,  etc.). 

This  species  is  probably  not  to  be  found  within  the  proper  bounds 
of  this  report.  It  is  introduced  chiefly  because  of  the  great  interest 
connected  with  the  remarkable  variability,  autotomy,  powers  of 
restitution  of  lost  rays,  or  even  of  the  entire  body,  from  a  part  of  one 
ray,  etc.,  which  have  been  so  well  and  carefully  studied  and 
described  by  Miss  Sarah  Monks,  in  the  works  cited  above. 

Other  species  of  the  genus  share  these  peculiarities,  for  example 
L.  guildingii,  which  I  have  studied  in  Bermuda. 

Miss  Monks,  op.  cit.,  1904  (pp.  596-600),  has  shown  that  this 
species  is  very  variable  in  number  of  rays,  madreporites,  etc.  The 
variations  in  number  of  rays  are  mostly  due  to  autotomy  and  irregu- 
lar restoration  of  rays,  so  that  there  may  be  from  five  to  eleven  rays. 
Though  five  is  the  normal  number,  regularly  six-rayed  specimens 
are  common. 

She  found  many  in  the  "  comet-form  "  and  succeeded  in  producing 
such  forms  by  cutting  off  rays  at  some  distance  from  the  disk. 
These,  in  the  course  of  six  months,  produced  a  "  comet-form  "  with 
a  new  disk. 


3IO  VERRILL 

Two  madreporites  are  common,  and  as  many  as  five  may  occur. 

Two  dorsal  pores  were  more  frequent  than  one,  in  the  pro- 
portion of  48  to  15 ;  eleven  had  three,  two  had  four,  and  one  had  five. 
The  genital  products  issued  from  various  points  along  the  rays. 

Its  range  is  from  Southern  California,  San  Pedro,  San  Diego,  etc., 
to  Panama  and  West  Colombia  and  the  Galapagos  Islands.  Very 
common  at  La  Paz,  Lower  California.  It  occurs  from  low  tide  to 
30  fathoms. 

Fisher  erroneously  quotes  "  L.  ornithopus"  Verrill,  1867,  as  a 
synonym  of  this  species.  No  such  determination  by  me  occurs  in 
the  work  cited  or  elsewhere.  On  page  344,  L.  ornithopus  is  given 
as  an  Atlantic  species,  "  probably  identical  "  with  L.  guildingU. 

Suborder  MYONOTA  Ludwig. 

Each  ray  has  a  pair  of  strong  internal  dorsal  muscles,  extending 
for  part  or  most  of  its  length.  Papular  areas  localized.  Pedicellariae 
usually  pectinate.  Podia  with  suckers. 

Family  BENTHOPECTINIDJE  (Verrill)  Fisher. 
Benthopectinida  and  Pontasterince  VERRILL,  Trans.  Conn.  Acad.  Sci.,  x,  pp.  200, 

217,  1899- 

Pararchasterida  SLADEN,  op.  cit.,  1889,  p.  4. 
Notomyofa  (order)  LUDWIG,  1910,  p.  435. 
Benthopectinidce  FISHER,  19116,  p.  120. 

An  almost  strictly  deep-sea  family.  Form  stellate;  disk  rather 
small;  rays  five,  elongated,  with  two  rows  of  thick,  spinose,  mar- 
ginal plates,  which  are  not  exactly  paired,  but  are  sometimes  alter- 
nate, or  nearly  so,  and  have  no  definite  fasciolated  grooves  between 
them ;  there  is  an  odd  interradial  marginal  plate  in  each  row,  in  some 
genera  (Benthopectininae).  Pedicellariae  are  pectinate  or  fascicu- 
late, rarely  bivalvular. 

Dorsal  abactinal  surface  is  covered  by  protopaxillse,  spinose  para- 
paxillae,  or  simple  spinose  plates,  rarely  with  true  paxillae. 

Papulae  may  be  distributed  over  most  of  the  lateral  surface  of  the 
rays,  or  may  be  limited  to  the  proximal  part  of  the  rays,  or  con- 
centrated in  specialized  areas  (papularia)  near  base  of  rays. 

Actinal  interradial  area  is  small  or  nearly  abortive ;  it  is  sometimes 
occupied  by  one  or  more  large  pectinate  pedicellariae. 

Similar  pectinate  pedicellariae  may  occur  between  the  plates,  on 
the  margins,  or  on  the  dorsal  surface.  Adambulacral  plates  are 
angular  and  have  elongated  furrow-spines  and  one  or  more  enlarged 
actinal  spines.  No  superambulacral  plates. 


SHALLOW- WATER   STARFISHES  311 

A  pair  of  muscle-bands  extends  along  the  medio-dorsal  region  for 
a  part  of  the  length  of  the  rays.  Dorsal  pore  usually  distinct.  Ambu- 
lacral  feet  have  suckers. 

Ludwig  (1910)  has  proposed  a  special  order  (Myonota)  for  this 
family,  on  account  of  the  presence  of  dorsal  radial  muscle-bands. 

It  seems  to  me  desirable  to  retain  two  subfamilies  in  this  group, 
though  recently  discovered  genera  render  them  less  distinct  than 
formerly. 

Subfamily  BENTHOPECTININJE  Verrill,  1894,  p.  245. 
Benthopectinida  (family)  VERRILL,  1899,  p.  217.    Ludwig,  1910,  p.  435. 

This  group  will  include  those  genera  in  which  an  odd  interradial 
marginal  plate  normally  exists  in  part  or  all  the  interradial  areas 
in  both  rows.  The  papulae  are  not  concentrated  in  specialized 
papularia,  though  they  may  be  confined,  in  some  genera,  to  the  basal 
part  of  the  rays.  Abactinal  plates  are  flat  or  low  tabulate,  usually 
lobed,  sometimes  stellate,  bearing  one  or  more  large,  sharp  spines 
and  usually  many  smaller  accessory  spines. 

This  is  a  strictly  deep-sea  group.  Several  species  have  been 
described  by  Dr.  Fisher  from  the  North  Pacific.  It  includes  the 
genera  Benthopecten  Verrill;  Myonotus  Fisher;  and  Nearchaster 
Fisher. 

Subfamily  PONTASTERIN&  Verrill,  1899. 

Cheirasteridce  (family)  LUDWIG,  1910,  p.  435. 
Benthopectinidee  (pars)  FISHER,  op.  cit,  1910,  p.  120. 

Odd  marginal  interradial  plates  are  not  normally  present.  Papulae 
are  confined  to  the  basal  parts  of  the  rays,  and  are  often  concentrated 
in  specialized  papularia. 

This  group  is  almost  entirely  confined  to  the  deep  sea.  Several 
species  are  known  from  the  North  Pacific  in  deep  water.  Only  one 
comes  within  my  bounds. 

Genus  Luidiaster  Studer. 

A  canthar chaster  VERRILL,  Proc.  U.  S.  Nat.  Mus.,  xvii,  p.  268,  1894. 
Luidiaster  STUDER,  1883,  p.  131.    Ludwig,  1910,  p.  451.    Fisher,  op.  cit,  19116, 
p.  127. 

Rays  usually  five,  angular,  tapered.  Disk  small.  Interradial 
actinal  plates  few,  confined  to  the  disk,  spinous.  Marginal  plates 
of  moderate  size,  more  or  less  alternate,  spiniferous;  those  of  the 
upper  series  smaller  than  those  of  the  lower,  rounded,  with  a  central 


312  VERRILL 

eminence  bearing  a  single  large,  movable  spine,  with  a  group  of 
small  spinules  around  its  base.  The  plates  of  the  lower  series  may 
bear  two  or  more  similar  large  spines  surrounded  by  spinules.  The 
upper  marginal  plates  form  a  narrow  margin  along  the  rays. 

The  dorsal  surface  is  covered  with  small,  unequal  plates  in  the 
form  of  protopaxillae  and  spinose  parapaxillae.1  The  latter  have  a  low, 
round  column  and  bear  a  large,  central,  articulated  spine  surrounded 
at  base  by  a  circle  of  small  spinules ;  they  are  found  on  the  disk  and 
along  the  median  part  of  the  rays.  The  protopaxillae  are  smaller,  and 
part  of  them  bear  only  small  spinules;  others  have  a  small  central 
spine.  The  papulae  occur  on  most  of  the  disk  and  the  entire  basal 
part  of  the  rays. 

Peculiar  double  pectinate  and  fascicled  pedicellariae  exist  on  the 
dorsal  surface  of  the  rays  and  disk,  and  a  single  one,  of  larger  size, 
occupies  the  center  of  each  actinal  interradial  area  (see  pi.  xxxiv, 
P,  P)  ;  in  one  case  a  similar  structure  replaces  the  two  upper  mar- 
ginal plates  in  the  interradial  angle.  These  large  actinal  compound 
pedicellariae  may  have  ten  to  twelve  incurved  papillae  on  each  side, 
while  those  of  the  dorsal  surface  have  usually  three  to  six.  Some  of 
the  latter  have  three  convergent  groups  of  curved  papillae.  The 
central,  dorsal  nephridial  pore  is  very  evident  and  is  surrounded 
by  papillae.  The  adambulacral  plates  have  a  salient  inner  angle,  and 
bear  a  horizontal  divergent  group  of  furrow-spines  and  a  transverse 
actinal  row  of  long  spines.  The  jaw-plates  are  large  and  bear  simple 
marginal  and  actinal  series  of  long  spines. 

LUIDIASTER  DAWSONI   (Verrill)   Ludwig. 
Plate  xxxin ;  plate  xxxiv;  plate  xxxv,  figure  2  (type). 

Archaster  dmvsoni  VERRILL,  in  Whiteaves,  Report  of  Progress  of  Geological 

Survey  of  Canada,  for  1878-79,  p.  1946  [p.  5],  1880. 
Acantharchaster  dawsoni  VERRILL,  Proc.  U.  S.  Nat  Mus.,  xvn,  p.  269,  1894. 

Fisher,  1910,  p.  549. 
Luidiaster  dawsoni  LUDWIG,   1910,  p.  452.     Fisher,   ignb,  p.   128,  pL  xxv, 

fig.  2;  pi.  xxvi,  fig.  3;  pi.  xxvn,  fig.  2;  pi.  LV,  figs.  3,  30;  pi.  LVI,  fig.  5;  pi. 

cxix,  fig.  2;  pi.  cxx. 

Rays  five,  long,  acute.  Radius  of  the  disk,  17  mm.;  of  rays, 
100  mm. ;  ratio,  1 : 6,  nearly.  The  rays  are  long,  flat,  regularly 
tapered.  The  upper  surface  is  loosely  covered  with  small  ossicles 
(protopaxillae),  those  toward  the  margins  of  the  rays  bearing  only 
circular  groups  of  very  minute  short  spinules ;  but  along  the  middle 

1  See  pages  280,  281,  for  explanations  of  these  terms. 


SHALLOW- WATER   STARFISHES  313 

region  of  the  rays  and  over  the  disk  they  are  spino-parapaxillae,  each 
bearing  a  long,  tapering,  acute  central  spine,  surrounded  at  base 
by  a  circle  of  small  spinules.  Between  the  plates  there  are,  over 
nearly  the  whole  surface,  numerous  papular  pores.  Along  each  ray, 
toward  the  marginal  plates,  there  are,  at  irregular  intervals,  singu- 
lar pectinate  pedicellariae,  consisting  of  groups  of  small  incurved 
spinules.  Usually  there  are  two,  three,  or  four  clusters.  Each 
cluster,  consisting  of  a  row  of  three  or  four  to  six  curved  spinules, 
form  one  group ;  and  the  ends  of  all  the  spinules  converge  to  a  pore 
in  the  center  of  the  group.  A  much  larger  elliptical  one,  composed  of 
ten  to  twelve  spinules  on  each  side,  occupies  each  actinal  interradial 
area. 

The  upper  marginal  plates  are  small,  but  prominent,  and  each 
bears  a  long,  rather  stout,  acute,  erect  spine,  surrounded  at  base  by 
a  group  of  slender,  unequal  spinules.  The  lower  marginal  plates 
mostly  bear  three  long  and  large  divergent  spines,  the  upper  one 
largest,  and  rather  longer  than  those  of  the  upper  plates;  between 
and  around  their  bases  there  are  slender  spinules.  The  adambulacral 
plates  bear  upon  the  inner  edge  a  rounded  group  of  about  six  very 
slender,  blunt  spines ;  the  two  lateral  ones  are  very  short ;  the  middle 
ones,  long;  outside  of  these  there  is  a  transverse  row,  usually  of 
three  much  longer  and  larger  blunt  spines. 

Dixon  Entrance,  British  Columbia,  n  fathoms  (type,  collected 
by  Dr.  G.  M.  Dawson,  1885). 

Dr.  Fisher  records  it  from  eleven  stations,  in  56  to  159  fathoms, 
from  Bering  Sea  to  Kadiak,  Alaska. 

Suborder  PAXILLOSA   (Perrier),  emended. 

Paxillosa  PERKIER  (as  an  order),  Exped.  Trav.  et  Talism.,  pp.  28,  29. 
Paxillosa  VERRILL,  Trans.  Conn.  Acad.,  x,  pp.  199,  200,  201,  1899. 

These  are  Phanerozona  in  which  the  two  rows  of  marginal  plates 
are  usually  well  developed  and  often  spinose,  and  usually  have  well 
developed  fasciolate  grooves  between  them.  The  dorsal  or  abactinal 
plates  are  usually  developed  in  the  form  of  true  columnar  paxillae 
or  spinopaxillae,  sometimes  as  pseudopaxillae.  Pedicellariae  are  often 
present,  usually  consisting  of  several  connivent  papillae,  or  spinule- 
like  structures,  surrounding  a  pore.  Sometimes  forceps-like  forms 
occur. 

Bivalve  valvular  pedicellariae  are  lacking.  Ambulacral  feet  are 
generally  pointed;  sometimes  flattened  and  natatory;  rarely  with  a 
small  terminal  knob,  but  without  a  sucker. 


314  VERRILL 

Ambulacral  ampullae  single  or  double.  Dorsal  pore  present  or 
absent.  Superambulacral  plates  usually  present;  sometimes  absent. 

The  Paxillosa  should  only  include  such  groups  as  have  neither  true 
bivalve  pedicellariae  nor  sucker-feet.  The  existence  of  true  paxilli- 
form  plates  on  the  dorsal  surface  cannot  be  made  an  invariable 
diagnostic  character,  for  they  occur  in  some  forms  of  Valvulosa. 
The  development  of  the  ambulacral  feet  varies  much  in  both  groups, 
and  depends  mainly  on  the  nature  of  the  bottom  commonly 
inhabited. 

The  families  represented  in  shallow  water  on  the  Northwest  coast 
are  Porcellanasteridae,  Astropectinidae,  and  Luidiidae.  See  families 
enumerated  on  page  283.  The  only  family  not  represented  is  Gonio- 
pectinidae,  a  deep-sea  group. 

Family  ASTROPECTINIDAE  Gray  (restricted). 

Astropectinidee    (pars)    GRAY,   Ann.    and    Mag.    Nat.    Hist.,   p.    140,    1840; 

Synopsis,  p.  2,  1866.     Sladen  (pars)  Voy.  Challenger,  xxx,  p.  174,  1889 

(includes  Luidia). 

Astropectinida  SLADEN,  op.  cit.,  1859,  p.  175. 
Astropectinida  (sense  extended)   FISHER,  op.  cit.,  191  ib,  p.  37.     (Analytical 

table  of  all  recognized  genera). 

Paxillosa  in  which  the  disk  is  usually  small  or  of  moderate  size, 
the  rays  often  much  elongated.  The  dorsal  surface  is  generally 
covered  with  highly  developed  true  paxillae  (rarely  with  para- 
paxillae  or  pseudopaxillae) ,  commonly  covering  fasciolated  inter- 
spaces with  intervening  simple  papulae. 

Both  rows  of  marginal  plates  are  usually  large,  the  inferomar- 
ginals  often  the  larger.  They  are  either  granulated  or  spinulose  and 
often  very  spinose,  with  more  or  less  simple  fasciolated  grooves 
between  them;  not  covered  by  a  thick  skin.  Adambulacral  plates 
usually  spinose  on  the  actinal  surface  and  with  a  divergent  row  of 
furrow-spines,  without  a  web. 

Pedicellariae  are  often  lacking ;  when  present  usually  f asciolate  or 
spiniform,  or  consisting  of  two  short  connivent  rows  of  spinules 
surrounding  a  special  pore.  Ambulacral  feet  in  two  rows,  large, 
usually  pointed,  never  with  suckers.  Ampullae  double.  Dorsal 
glands  and  pore  usually  present.  Superomarginal  plates  present. 
Interactinal  plates  wanting,  or  more  or  less  numerous  in  regular 
simple  rows,  usually  spinulose  and  with  fasciolated  grooves  between 
all  the  rows,  but  without  marginal  webs. 

The  aproctous  condition,  formerly  supposed  to  be  characteristic 
of  the  family,  is  unreliable,  for  in  nearly  all  the  genera  referred  to  it 


SHALLOW- WATER  STARFISHES  31$ 

by  Mr.  Sladen  there  is  a  perfectly  well  defined  dorsal  or  "  anal  "  pore, 
and  in  some  of  the  genera  the  pore  is  even  elevated  on  the  summit 
of  a  dorsal  cone  or  chimney  (Psilaster,  I ly aster,  etc.).  This  pore, 
which  I  have  designated  as  "  p sen danu s "  or  "nephridial  pore," 
serves  in  each  of  these  families  (and  in  Asterioidea  generally)  mainly 
for  the  discharge  of  the  secretion  of  the  branched,  dorsal,  glandular 
organs,  probably  nephridial  in  function,  situated  above  the  stomach 
and  often  called  "  ccecal  appendages."  These  glands  are  sometimes 
absent.  Whether  the  central  pore  serves  as  a  true  anus  in  any  of 
these  starfishes  is  very  doubtful,  for  the  intestine  is  usually  nearly 
or  quite  abortive.  In  any  case,  it  is  often  impossible  to  ascertain 
this  point  without  actual  dissection  of  alcoholic  or  fresh  specimens, 
which  are  often  not  available. 

I  have  been  unable  to  demonstrate  a  functional  intestine  in  any 
case,  and  have  never  observed  fcecal  matter  in  the  so-called  intestine ; 
nor  have  I  ever  seen  such  substances  discharged  by  the  dorsal  pore 
by  any  living  starfish,  though  I  have  studied  thousands  of  living 
specimens,  including  numerous  genera  and  families.  In  Pterasteridae, 
however,  the  intestine  is  very  obvious. 

The  existence  of  definite  fascicles  of  specialized  spinules  or 
papillae  on  the  margins  of  the  plates,  so  as  to  form  covered  chan- 
nels along  their  sutures,  in  this  and  the  related  families,  is  evidently 
a  character  both  of  morphological  and  of  physiological  importance. 
The  existence  of  such  fascioles  is  correlated  with  the  mode  of  life. 
Such  forms  as  have  them  appear  commonly  to  live  more  or  less 
buried  in  soft  mud  or  sand,  and  the  fascioles  are  evidently  for  the 
purpose  of  providing  a  free  circulation  of  water  around  the  whole 
surface  of  the  body,  both  to  provide  for  respiration  and  to  keep  the 
surface  of  the  body  free  from  dirt.  The  paxilliform  plates  and 
spinules  also  contribute  to  both  these  functions  and  also  serve  to  pro- 
tect the  dermis  and  papulae  from  dirt. 

The  typical  Astropectinidae  are  among  those  best  provided  with 
fascioles  and  with  the  most  highly  developed  forms  of  paxillae.  They 
are  also  those  that  are  eminently  dwellers  on  and  beneath  mud  and 
sand.  The  pointed  form  of  the  ambulacral  feet  is  correlated  with 
the  same  habit,  but  they  are  not  confined  to  such  places. 

The  genera  Ctenodiscus,  Porcellanaster,  Luidia,  and  allies  have 
similar  but  even  more  specialized,  structural  adaptations  for  the  same 
purposes. 

Dr.  Fisher  thinks  that  it  is  not  possible  to  divide  the  family  into 
subfamilies,  as  others  have  done.  Owing  to  the  recent  discovery,  in 


VERRILL 

the  deep  sea,  of  numerous  more  or  less  intermediate  genera,  the 
limits  of  such  groups,  as  well  as  those  of  recognized  families  and 
genera,  have  become  more  and  more  indefinite.  Probably  future 
discoveries  will  make  them  even  more  so. 

Still  it  is  convenient  to  recognize  subfamily  groups,  even  if  not 
always  closely  definable  by  any  one  or  two  characters.  It  is  the 
combination  of  several  characters  that  counts  most  in  all  the  larger 
groups. 

ANALYTICAL  TABLE  OF  NORTHWEST  AMERICAN   SHALLOW- 
WATER  GENERA  OF  ASTROPECTINID^. 

Marginal  plates  thick,  convex,  spinulose  on  the  whole  surface;  separated 
by  deep  sutural  grooves,  which  are  usually  bordered  by  many  rows 
of  fine  fasciolated  spinules.  Interactinal  plates,  when  present, 
convex,  spinulose,  pseudopaxilliform,  and  fasciolated. 

A.  Marginal  plates  of  both  series  large;  lower  ones  mostly  much  prolonged, 

reaching    the    adambulacrals.     Dorsal    plates    are    true    columnar 
paxillae;  interactinal  plates  few  or  none.    Rays  elongated. 
Astropecten  (restricted). 

AA.  Lower  marginal  plates  are  not  so  much  prolonged  adorally.  Inter- 
actinal plates  are  in  rows,  forming  a  triangular  area. 

B.  Dorsal  plates  form  convex  pseudopaxillae  with  stellate  bases ;  interactinal 

plates  are  flat  or  convex  pseudopaxillse,  in  three  or  more  paired 
rows,   with    an    odd   interradial   row;    not    evidently   imbricated; 
lower  marginal  plates  are  about  equal  in  length  to  two  adambula- 
crals; spinose  centrally.     Rays  rather  long.     Marginal  fasciolated 
grooves  are  simple  and  narrow.     Adoral  marginal  adambulacral 
spines  are  deep  in  the  grooves. 
Bunodaster.     Type,  B.  ritteri  Verrill. 
BB.  Dorsal  plates  are  true  columnar  paxillas  or  parapaxillae. 

C.  Lower  marginal  plates  are  usually  spinose  and  spinulose,  as  in  Astropecten. 

Rays  elongated. 

Astropectinides  Verrill.    Type,  A.  mesacutus  (Sladen).    Antarctic. 
CC.  Marginal  plates  not  spinose,  but  covered  with  minute  spinules;  sutural 
grooves  wide  and  deep.    Rays  short  or  rather  short.     Odd  actinal 
interradial  plates  are  present. 

D.  Marginal  plates  short  and  about  equal  in  number  to  the  adambulacral 

plates ;    upper    ones   much   smaller    than   the   lower ;    both   series 
become   oblique   distally,   the   sutures    slanting  adorally.      Actinal 
plates  have  a  raised  center,  or  carina,  and  strong  lateral  fascioles. 
Leptychaster  Smith.    Type,  L.  kerguelenensis. 

DD.  Marginal  plates  few,  large,   subrectangular,   subequal   in  breadth,  each 

about  as  long  as  two  adambulacrals;  grooves  wide,  not  notably 

oblique  distally.     Interactinal  plates  fasciolated,  in  three  or  more 

rows,  rounded,  convex,  tabulate,  not  carinate ;  a  few  unpaired  ones. 

Glyphaster.     Type,  G.  anomalus  (Fisher). 


SHALLOW-WATER   STARFISHES  317 

Genus  Astropecten  Gray  (emended). 

Astropecten  (pars)  GRAY,  Ann.  and  Mag.  Nat.  Hist.,  vn,  p.  180,  1840;  Sy»- 
tem,  p.  3,  1866.  Muller  and  Troschel,  Syst.  Aster.,  p.  67,  1842.  Siaden. 
op.  cit.,  1889,  p.  193.  Fisher,  op.  cit.,  191  ib,  p.  55. 

Stellaria  NARDO,  Oken's  Isis,  p.  716,  1834  («on  Muller,  Mollusca,  1832). 

Asterias  L.  AGASSIZ,  Prod.,  p.  191,  1835. 

Rays  more  or  less  elongated,  subacute,  flat  above,  with  true  abac- 
tinal,  stellate  paxillae,  and  stout,  fasciolated  marginal  plates.     The 
upper  marginals  are  convex,  encroaching  more  or  less  on  the  upper 
side  of  the  disk,  usually  spinulose,  sometimes  almost  granulose,  and 
often  with  one  or  two  superior  spines  or  tubercles.    Lower  marginals 
spinulose  and  spinose,  prolonged  actinally,  most  of  them  reaching 
the  adambulacral  plates;  but  one  or  two  interradial  pairs  may  not 
reach  the  adambulacral  plates,  and  in  that  case  one  or  two  pairs  of 
small  adoral  interactinal  ossicles  may  intervene,  but  these  do  not  form 
regular  rows,  nor  an  important  triangular  area.    The  species  having 
an  important  group  of  actinal  plates  are  here  separated  as  a  genus, 
Astropectinides.     The  lower  marginals  are  closely  spinulose  and 
fasciolated  laterally.     Adambulacral  plates  about  equal  in  number 
to  lower  marginals,  the  inner  end  angular.    They  have  a  horizontal 
group  of  about  three  furrow-spines  and  a  divergent  stellate  group 
on  the  actinal  side,  often  with  the  central  spine  enlarged.     The 
papulae  stand  singly  around  the  dorsal  paxillae,  generally  absent  along 
the  median  line.     Dorsal  pore  usually  lacking,  sometimes  present. 
Superambulacral    plates    well    developed.      Pedicellariae    generally 
lacking. 

ASTROPECTEN  SIDEREALIS  Verrill,  sp.  nov. 
Plate  L,  figure  6  (details). 

Astropecten  armatus  FISHER  (non  Gray),  op.  cit.,  igiib,  p.  56,  pi.  r,  figs,  i,  2; 
pi.  vu,  figs.  3,  6;  pi.  L,  fig.  4;  pi.  LI,  fig.  3. 

A  variable  species.  The  type  has  the  following  characters :  Disk 
of  moderate  size,  thick;  rays  five,  rapidly  tapered,  acute.  Radii, 
14  mm.  and  58  mm. ;  ratio,  1 : 4.15. 

The  upper  marginal  plates  are  short,  convex,  prominent,  their 
upper  ends  rising  above  the  level  of  the  abactinal  paxillae.  They  are 
separated  by  rather  wide  and  deep  fasciolated  grooves ;  those  in  the 
interradial  angles  are  externally  wedge-shaped,  with  the  sutures 
vertical,  but  distally  the  sutures  become  oblique  and  the  plates 
scarcely  wider  than  long.  Most  of  their  outer  surface  is  closely 
covered  with  small,  rounded  or  hexagonal  granules,  which  grade 


3l8  VERRILL 

into  the  small  spinules  of  the  fascicles  on  the  sides.  At  the  summit 
of  more  or  less  of  the  proximal  plates  there  is  a  small,  nipple-shaped 
or  thimble-shaped  spine,  which  is  liable  to  be  broken  off,  leaving  a 
small,  pit-like  scar. 

The  dorsal  paxillae  bear  small,  regular,  stellate  groups  of  short, 
somewhat  unequal,  clavate  or  capitate  spinules.  In  the  larger  ones 
there  is  usually  a  central,  larger,  capitate  spinule,  surrounded  by  a 
circle  of  about  six  to  eight  smaller  ones ;  these  are  surrounded  by  a 
marginal  circle  of  about  ten  to  fifteen  of  about  the  same  size. 

The  lower  marginal  plates  are  strongly  spinose ;  most  of  the  sur- 
face is  covered  with  small,  flat,  more  or  less  imbricated  spinules. 
There  is  a  submedian  row  of  six  to  eight  unequal,  roundish,  acute 
spines,  increasing  in  size  outwardly,  the  last,  or  next  to  the  last,  being 
much  the  largest,  stout  and  acute,  not  much  flattened;  sometimes 
there  is  a  smaller  slender  one  above  the  largest. 

Adambulacral  plates  have  a  convex  inner  or  furrow-group  of 
about  four  slender  spines,  of  which  the  innermost  is  larger,  longer, 
angular,  and  stands  in  front  of  the  others.  On  the  actinal  side  there 
is  a  central,  erect,  unusually  large,  flat,  blunt  or  truncate,  often 
grooved  spine,  and  back  of  this  a  row  of  three  or  four  much  smaller 
spines,  of  similar  shape,  on  the  outer  and  proximal  margins  of  the 
plate. 

The  adoral  and  epioral  spines  are  very  numerous,  short,  stout, 
obtuse. 

It  varies  greatly  in  the  length  and  sturdiness  of  the  rays,  in  the  size 
of  the  superomarginal  plates,  and  especially  in  the  number  and  form 
of  the  tubercles  or  spines  on  the  superomarginal  plates.  These  may 
be  reduced  to  a  single  imperfect  series  of  small  tubercles,  or  there 
may  be  one  continuous  series,  or  two  imperfect  series — rarely  two 
nearly  complete  series.  They  may  be  lacking  proximally  and  pres- 
ent distally,  or  vice  versa.  Sometimes  they  become  spine-like,  espe- 
cially the  outer  ones ;  but  more  often  they  are  short,  stubby  cones  or 
blunt  tubercles.  Other  parts  also  vary.  Generally,  however,  the 
enlargement  of  the  central  spinules  of  some  of  the  paxillae,  the 
differentiation  of  the  paxillae  along  the  median  line,  and  especially 
the  scale-like,  flattened,  inferomarginal  spinelets,  remain  pretty 
constant. 

This  species,  in  form  and  general  appearance,  resembles  A.  aer- 
stedii  Liitken,  of  Central  America,  which  is  now  generally  considered 
identical  with  the  true  armatus  of  Gray.  This  resemblance  is  due 
in  large  part  to  the  character  of  the  marginal  spines  in  the  interradial 


SHALLOW- WATER   STARFISHES 

regions,  and  the  size  and  form  of  the  marginal  plates.  The  easiest 
distinctive  mark  is,  perhaps,  the  flattened  and  imbricated  condition 
of  the  inferomarginal  spinules. 

I  have  compared  it  with  a  type-specimen  of  oerstedii,  received 
from  Dr.  Liitken  himself.  The  upper  marginal  plates  in  the  latter 
have  two  small  spines ;  the  dorsal  paxillary  spinules  are  longer ;  the 
lower  marginal  plates  have  two  distinct  rows  of  larger  spines,  and 
the  small  spinules  are  more  slender  and  not  imbricated,  nor  flat  and 
scale-like;  the  adambulacral  spines  are  similar,  but  more  slender, 
especially  the  larger  central  spine  of  the  actinal  side,  which  is,  also, 
less  flattened. 

This  species  is  allied  to  A.  calif  ornicus,  but  has  coarser  paxillary 
spinules,  with  a  larger  central  one,  and  different  spinules  on  the 
marginal  and  adambulacral  spines.  The  latter  never  has  spines  or 
tubercles  on  the  upper  marginal  plates. 

Near  San  Francisco  (Prof.  W.  E.  Ritter).  Type  in  Yale  Museum. 

Professor  Fisher  (op.  cit.,  19116,  p.  60)  has  given  a  number  of 
localities  for  this  species,  under  the  name  A.  armatus.  They  range 
from  off  San  Pedro  and  Long  Beach,  California,  to  San  Diego,  in 
4  to  130  fathoms.  He  considers  that  its  range  extends  to  Panama 
and  Ecuador  (as  armatus),  but  I  consider  the  southern  form  distinct. 

VARIATIONS. 

Professor  Fisher  has  given  a  detailed  description  and  several 
figures  of  this  species,  and  has  very  fully  discussed  its  variations. 
Therefore  it  is  not  necessary  to  give  many  details  here.  The  above 
description  was  written  in  1904,  long  before  his  paper  appeared, 
and  was  based  on  a  rather  average  specimen,  as  the  type. 

ASTROPECTEN  CALIFORNICUS  Fisher. 

Plate  L,  figure  5  (details) ;  plate  c,  figure  i;  plate  ci,  figures  i,  2;  plate  CH, 

figures  i,  2. 

Astropecten  calif  ornicus  FISHER,  op.  cit,  1906,  p.  299;  op.  cit,  IQII&,  p.  61, 
pi.  vi,  figs,  i,  2;  pi.  vii,  fig.  i ;  pi.  L,  fig.  5 ;  pi.  LI,  figs.  2,  20. 

Disk  small,  but  rather  thick ;  rays  five,  long,  rather  narrow,  regu- 
larly tapered,  acute.  Radii,  17  mm.  and  77  mm. ;  ratio,  i :  4.5. 

Upper  marginal  plates  numerous,  not  very  wide,  nor  much 
encroaching  on  the  dorsal  surface,  convex,  separated  by  deep  fascio- 
late  grooves,  without  special  spines,  but  covered  with  unequal  small 
spinules,  of  which  those  in  two  or  three  rows  along  the  middle  are  a 
little  larger  and  capitate  or  like  rounded  granules,  while  the  rest  are 
longer,  slender  and  blunt. 


320  VERRILL 

The  dorsal  paxillae  are  small,  even,  and  much  crowded,  with 
regularly  stellate,  small,  short,  clavate  spinules,  of  which  there  may 
be  ten  to  fourteen  marginal  and  five  to  eight  central  ones  on  the 
larger  paxillge.  On  the  center  of  the  disk  the  paxillae  become  smaller 
and  very  much  crowded,  so  that  they  seem  to  blend  together.  On 
the  bases  of  the  rays  they  are  about  i  mm.  to  1.2  mm.  broad. 

The  inferomarginal  plates  are  large  and  very  spinose ;  they  bear 
two  submedian  rows  of  acute  spines,  increasing  in  size  upward,  the 
upper  two  or  three  being  much  larger  and  longer  than  the  rest,  and 
on  the  basal  plates  they  rapidly  increase  in  size  and  become  flattened, 
somewhat  paddle-shaped,  with  acuminate  tips.  The  rest  of  the  sur- 
face is  covered  with  numerous  small,  somewhat  imbricated  and  flat- 
tened, blunt  spinules. 

The  adambulacral  plates  have  a  furrow-series  of  three  or  four 
rather  long,  slender,  divergent  spines;  and  on  the  actinal  side  four, 
or  sometimes  five,  much  larger,  flattened  or  paddle-shaped,  obtuse, 
erect  spines,  usually  standing  two  by  two.  The  central  spine  of  the 
furrow-series  is  stouter  than  the  others,  but  stands  a  little  farther 
within  the  groove. 

No  pedicellariae  were  found. 

Off  San  Francisco  (Prof.  W.  E.  Ritter)  ;  Pacific  Grove  (Prof. 
W.  R.  Coe) .  This  species  has  a  known  range  from  Monterey  Bay 
to  Lower  California. 

Dr.  Fisher  records  it  from  numerous  stations,  from  off  Monterey 
to  Guadalupe  Island,  in  10  to  244  fathoms.  Abundant  off  Monterey ; 
off  San  Pedro ;  off  Santa  Barbara,  etc.,  in  25  to  50  fathoms. 

ASTROPECTEN  ORNATISSIMUS  Fisher. 

Astropecten  ornatissimus  FISHER,  op.  cit,  19066,  p.  119;  op.  cit.,  19116,  p.  67, 
pL  vi,  figs.  3,  4;  pi.  vn,  fig.  2;  pi.  LI,  figs.  i-ic. 

Allied  to  A.  calif ornicus.  Differs  in  having  larger  dorsal  paxillae, 
with  longer,  more  slender,  and  more  spaced  spinules ;  in  having  the 
paxillae  more  uniform  in  size  across  the  ray,  not  more  elongated 
medially,  nor  smaller  over  a  large  central  area,  of  the  disk ;  in  having 
more  slender,  longer  and  more  tapered  adambulacral  spines;  in 
having  the  upper  edge  of  each  ambulacral  plate  produced  into  a  thin 
lamina  between  the  pairs  of  ampullae,  with  its  edge  serrulate. 

Greater  radius,  56  mm. ;  lesser,  14  mm. ;  ratios,  i :  4  (Fisher). 

I  have  not  received  this  species.  Mr.  Fisher  records  it  from 
sixteen  stations  from  off  San  Pedro  and  Catalina  Island,  to  Guada- 
lupe and  Cerros  Islands,  Lower  California,  in  47  to  207  fathoms. 


SHALLOW-WATER   STARFISHES  321 

Genus  Astropectinides  Verrill,  nov. 
Type,  A.  mesacutus  (Sladen),  as  Astropecten. 

This  genus  is  proposed  for  those  species,  usually  referred  to 
Astropecten,  which  have  a  well  developed  triangular  actinal  area, 
covered  with  two  or  more  rows  of  interactinal  plates  parallel  with 
the  adambulacrals  and  corresponding  to  them  in  radial  length.  The 
proximal  inferomarginal  plates  are  correspondingly  abbreviated, 
not  prolonged  nearly  or  quite  to  the  jaws,  as  in  Astropecten. 
The  dorsal  paxillae  and  upper  marginal  plates  are  as  in  Astropecten. 

Several  species,  described  as  Astropecten,  besides  the  type,  belong 
to  this  genus.  Among  them  are  A.  callistus  (Fisher)  and  A.  cteno- 
phorus  (Fisher),  Hawaiian  Islands.  The  genus  is  extralimital. 

The  type,  A.  mesacutus  Sladen  (1883;  and  op.  cit.,  1889,  p.  219, 
pi.  xxxiv,  figs.  5,  6 ;  pi.  xxxvm,  figs.  7-9),  has  twenty  to  thirty  spinu- 
lose  plates  in  each  interradial  area.  It  is  from  the  Antarctic  Ocean 
and  the  South  Atlantic,  in  44  to  90  fathoms. 

Genus  Bunodaster  Verrill. 
Plate  LXXXVI,  figures  i,  10;  plate  civ,  figures  i,  10;  plate  cv,  figures  i,  10. 

Bunodaster  VERRILL,  Amer.  Naturalist,  xun,  p.  554,  fig.  4,  September,  1909. 
Fisher,  op.  cit.,  19110,  p.  164;  ignb,  p.  40. 

Superficial  appearance  much  as  in  some  dorsally  unarmed  species 
of  Astropecten,  but  with  smaller  marginal  plates. 

Form  stellate  with  elongated  rays.  Dorsal  surface  covered  with 
low,  convex,  somewhat  hemispherical  plates,  which  are  of  the  nature 
of  parapaxillae,  with  enlarged,  somewhat  lobed  or  stellate  concealed 
bases,  the  lobes  slender  and  articulated,  with  single  papulae  between 
them.  The  surface  of  the  outer  boss,  or  raised  part,  is  covered  with 
numerous  small,  slender,  divergent,  spinules,  which  form  simple,  deep 
fascicles  at  the  margins. 

Marginal  plates  not  very  thick,  convex,  all  closely  spinulose  and 
the  lower  ones  spinose ;  the  upper  ones  are  rather  small,  in  the  inter- 
radial  arcs,  where  the  lower  ones  are  not  much  prolonged  adorally, 
but  end  abruptly  against  a  triangular  area  of  fascicled  and  spinulose 
actinal  plates,  which  are  flattish  or  convex  and  form  rows  parallel 
with  the  adambulacrals,  or  three  V-shaped  rows  with  two  odd  inter- 
radial  plates  in  line.  The  adambulacral  plates  are  the  same  in  length 
as  the  synactinals ;  they  have  a  horizontal  furrow-group  of  three  or 
four  spines  and  an  actinal  divergent  group  of  similar  ones. 

22 


322  VERRILL 

Fasciolated  grooves  extend  between  the  adambulacral  and  most  of 
the  interactinal  plates.  Jaws  are  very  prominent  on  the  actinal  sur- 
face and  bear  numerous  peroral  and  adoral  spines  on  the  sides  and 
summit.  The  lateral  peroral  spinules  are  fasciolated;  the  actinal 
groups  or  rows  are  connivent. 

This  genus  is  undoubtedly  closely  allied  to  Blakiaster,  Persephon- 
aster  and  Leptycliaster,  especially  to  the  two  former. 

Dr.  Fisher  thinks  (op.  cit.,  191  ib,  p.  40)  that  it  ought  to  be  united 
with  Blakiaster,  which  he  carefully  describes  from  the  type.  I  have 
also  studied  the  same  type.  It  differs  from  that  genus,  however,  in 
the  less  massive  and  differently  shaped  marginal  plates,  which  are 
less  complex  and  have  more  fully  developed  fascioles.  The  lower 
marginals  have  the  spines  in  a  central  row  or  group,  not  submar- 
ginal.  There  is  no  differentiated  median  radial  series  of  dorsal 
ossicles ;  nor  any  notable  number  of  secondary  ossicles,  so  evident  in 
Blakiaster;  nor  any  median  radial  area  destitute  of  papulae,  as  in  the 
latter.  There  are  well  defined  fascioles  between  the  adambulacral 
and  interactinal  plates,  not  found  in  the  latter ;  and  the  adoral  adam- 
bulacral spines  are  deeply  sunken  in  the  groove,  as  in  Persephon- 
aster,  but  not  in  Blakiaster  (type,  B.  conicus  Perrier). 

The  abactinal  plates  are  thinner,  more  regularly  stellate,  and  more 
delicately  articulated  by  the  slender  radial  lobes  than  in  the  latter, 
so  that  the  test  is  more  flexible.  In  Blakiaster  the  lobing  is  irregular 
and  the  edges  overlap. 

In  many  of  these  characters  it  is  nearer  to  Persephonaster,  but 
the  latter  lacks  the  odd  interradial  interactinal  plates,  found  in  this 
genus  and  in  Leptychaster. 

From  the  latter  it  differs  in  the  characters  of  the  abactinal 
ossicles,  in  the  less  profoundly  fasciolated  grooves  between  the 
marginal  plates,  and  in  other  characters ;  but  it  is  perhaps  as  nearly 
allied  to  the  latter  as  to  Blakiaster. 

It  is,  in  fact,  a  sort  of  connecting-link  between  these  three  genera. 
Possibly  it  may  eventually  be  desirable  to  unite  it  with  Leptychaster 
and  Blakiaster  in  one  larger  genus,  should  more  connecting  species 
be  discovered. 

BUNODASTER  RITTERI  Verrill. 

Plate  LXXXVI,  figures  i,  10  (details)  ;  plate  civ,  figures  i,  2  (type)  ;  plate  cv, 

figures  i,  10  (type,  enlarged)  ;  text-figures  Nos.  15,  16. 
Bunodaster  ritteri  VERRILL,  op.  cit.,  XLIII,  p.  554,  fig.  4,  1909. 

Rays  five,  regularly  and  rapidly  tapered,  slender,  subacute;  neatly 
stellate.  Disk  rather  thin,  flat,  about  as  wide  as  the  length  of  the 


SHALLOW-WATER   STARFISHES 


323 


rays,  with  broadly  curved  interradial  margins.     Radii,  10  mm.  and 
32  mm. ;  ratio,  i :  3.2. 

Upper  marginal  plates  regular,  not  very  large,  a  little  convex,  nar- 
row and  encroaching  very  little  on  the  dorsal  side  in  the  interradial 
angles,  but  on  the  rays  becoming  more  conspicuous,  as  seen  from 
above,  slightly  oblique,  squarish.  The  two  rows  at  the  middle  of  the 
ray  are  separated  by  a  space  only  about  equal  to  the  width  of  a  plate ; 
they  are  regularly  and  uniformly  and  closely  covered  by  small,  erect, 
divergent,  blunt,  rough  spinules,  and  separated  by  narrow,  deep, 
finely  fasciolated  grooves. 


FIG.  15. 

Bunodaster  ritieri  Verrill,  type.  Portion  of  under  side  of  a  ray;  ad,  adambulacral  spines; 
o.  a',  two  rows  of  interactinal  plates  and  spines;  m,  m,  two  infer  omarginal  plates  and  spines 
enlarged. 


FIG.  16. 

Bunodaster  ritteri  Ver.,  type.    A  group  of  dorsal  plates  and  papular  pores  from  middle  of 
the  base  of  a  ray,  seen  from  the  inside,  much  enlarged. 

The  dorsal  surface  is  covered  with  rather  large,  round,  mostly 
equal,  convex,  stellate  paxillae,  each  covered  with  a  large  number  of 
regularly  divergent,  small,  nearly  equal,  obtuse,  rough  spinules,  of 
which  the  larger  have  from  twenty  to  thirty.  Those  at  the  margins 
form  simple,  deep  fascicles.  When  the  spinules  are  removed,  the 
plates  are  in  the  form  of  low,  round-topped  or  nearly  hemispherical 
bosses,  mostly  nearly  equal  in  size  and  form,  with  only  a  few  smaller 
ones,  here  and  there,  and  separated  by  intervals  of  flexible  integu- 
ment, with  about  six  papulae  standing  singly  and  regularly  around 
each  plate.  The  papulae  are  lacking  on  a  small  central  area  of  the 


324  VERRILL 

disk  and  on  a  small  triangular  interradial  area  close  to  the  marginals, 
where  the  ossicles  are  more  closely  joined.  They  occur  regularly 
on  the  rest  of  the  disk  and  on  the  entire  breadth  of  the  rays,  except 
distally,  near  the  tips.  There  is  no  median  radial  band  of  ossicles 
differing  in  size  or  form  from  those  adjacent  and  not  surrounded 
by  papulae,  as  is  the  case  in  Blakiaster. 

Seen  from  the  inner  side,  the  dorsal  ossicles  are  all  alike,  except 
for  a  few  rather  smaller  ones,  and  lie  in  one  plane.  Their  inner  or 
basal  portion  is  roundish,  mostly  with  six  slender,  radiating  lobes; 
or  they  may  be  called  six-rayed  stellate.  They  are  articulated  only  by 
the  slender  ends  of  these  radial  lobes,  which  ordinarily  do  not  show 
at  all  on  the  outer  surface.  The  papular  pores  are  between  the  radial 
lobes. 

The  inferomarginal  plates  are  short  in  the  radial  direction,  in  the 
interradial  arcs,  scarcely  exceeding  the  upper  ones,  wedge-shaped, 
thick  below.  Towards  the  middle  of  the  rays  they  become  a  little 
oblique,  nearly  as  wide  as  long,  somewhat  rhombic,  near  the  tips  of 
the  rays  they  become  small,  apple-seed  shaped,  and  imbricated 
obliquely.  They  are  covered  with  small  spinules ;  like  the  upper  ones, 
and  mostly  bear  a  small  cluster,  or  short  median  row,  of  three  or 
four  small,  slender,  acute  spines,  much  longer  than  the  spinules. 

The  triangular  interradial  areas,  below,  are  flattened,  with  a 
pavement  of  slightly  convex,  roundish  plates,  mostly  in  three  pairs 
of  divergent  rows  parallel  with  the  ambulacra;  they  are  covered 
with  stellate  groups  of  numerous  spinules,  like  those  of  the  marginal 
plates,  but  rather  longer.  An  odd  interradial  plate  lies  between  the 
ends  of  the  second  and  third  rows. 

The  adambulacral  plates  have  a  convex  marginal  series  of  about 
four  rather  long,  slender,  divergent  spines ;  and  one  actinal  divergent 
or  stellate  group  of  about  six  to  eight  smaller  and  more  slender 
spines. 

Epioral  spines  small,  slender,  and  very  numerous,  in  two  large 
series.  Perorals  a  little  stouter. 

Apical  oral  spines  four,  rather  stout,  high  up  on  the  jaws  and 
nearly  concealed  by  the  adjacent  epiorals.  The  adoral  adambulacral 
spines  are  deep  within  the  grooves,  opposite  the  large  and  promi- 
nent jaw-plates,  on  the  edges  of  the  strongly  compressed  adoral 
plates,  forming  a  nearly  simple  curved  row. 

When  the  epioral  spines  are  removed,  the  two  prominent  ridges  of 
the  actinal  face  of  the  jaw-plates  are  closely  united,  forming  a  con- 
spicuous, elevated,  narrowly  elliptical  carina. 


SHALLOW-WATER   STARFISHES  325 

The  terminal  or  ocular  plate  of  the  rays  is  relatively  large,  thick, 
trapezoidal,  longer  than  wide.  It  is  reached  by  about  two  or  three 
rows  of  very  small  dorsal  plates. 

No  pedicellariae  were  found. 

Off  San  Francisco  (Prof.  W.  E.  Ritter).    Only  one  specimen  seen. 

Genus  Leptychaster  Smith. 

Leptychaster  SMITH,  Ann.  and  Mag.  Nat.  Hist,  ser.  4,  xvn,  p.  no,  1876. 
Leptoptychaster  SMITH,  Phil.  Trans.  Roy.  Soc.,  Zool.,  CLXVIII,  p.  278,  1879. 
/  Parastropecten  LUDWIG,  Mem.  Mus.  Comp.  Zool.,  xxxn,  p.  76,  1905. 
Leptychaster  (pars)  FISHER,  op.  cit.,  191  ib,  p.  42. 

Stellate,  with  five  short  rays,  rarely  six ;  disk  broad. 

Marginal  plates,  thick,  oblique,  at  least  distally,  the  upper  ones 
often  smaller  than  the  lower  and  not  encroaching  much  on  the  abac- 
tinal  side,  proximally;  prominent  or  carinate  in  the  middle,  with 
wide  and  deep  fasciolated  grooves  between  them ;  covered  with  small 
spinules,  which  become  larger  in  the  grooves.  Dorsal  plates  are  true 
paxillse,  with  roundish  or  somewhat  stellate  bases,  in  the  papular 
areas ;  their  spinelets  are  usually  very  small  and  numerous. 

The  interactinal  plates  form  an  important  triangular  area.  They 
are  spinulose,  thick  or  carinate,  with  deep  fasciolated  grooves 
between  them.  They  are  arranged  in  transverse  rows  running  to  the 
adambulacral  plates ;  or  they  may  be  said  to  be  in  rows  parallel  with 
the  ambulacra,  with  a  few  unpaired  interradial  plates.  The  trans- 
verse rows  of  interactinals  nearly  correspond  in  number  to  the  mar- 
ginals, except  in  the  middle  of  the  interradial  areas,  where  there  are 
more.  The  adambulacral  plates  have  a  group  of  slender  furrow- 
spines  and  a  divergent  group  of  similar  ones  on  the  actinal  side. 

Dr.  Fisher  thinks  that  Parastropecten  Ludwig  and  Glyphaster 
Verrill  should  be  united  with  Leptychaster. 

The  discovery  of  L.  pacificus  and  other  somewhat  intermediate 
forms  renders  this  view  somewhat  reasonable.  It  would  seem,  how- 
ever, that  Glyphaster  can  be  retained  at  least  as  a  subgenus,  if  not  a 
genus,  for  those  species  which,  like  anomalus,  have  robust  and 
squarish  upper  marginal  plates,  and  the  abactinal  plates  of  the 
papular  areas  only  slightly  lobed.  The  inferomarginal  plates  are 
not  oblique  in  the  latter,  and  each  corresponds  to  two  rows  of  inter- 
actinals instead  of  one. 

Whether  Parastropecten  Lud.  is  strictly  synonymous  with  Glyph- 
aster  is  uncertain.  The  type  is  P.  inermis  Lud. 

The  type  of  Leptychaster  is  L.  kerguelenensis  Smith,  from  the 
Antarctic.  In  a  later  paper  he  used  a  revised  spelling  of  the  word 


326  VERRILL 

(Leptoptychaster) ,  which  has  been  adopted  by  Sladen  and  many 
other  writers.    The  change  seems  to  be  uncalled  for  and  pedantic. 

The  type  species  is  known  to  carry  its  eggs  and  young  on  the  back, 
under  the  paxillary  spines.  This  has  not  been  observed  in  either  of 
the  northern  species. 

LEPTYCHASTER  ARCTICUS  (Sars)  Sladen. 

Astropecten  arcticus  M.  SARS,  Reise  Lofoden  og  Finmarken,  Nyt  Mag.  Nat., 

vi,  p.  161,  1851. 

Archaster  arcticus  VERRILL,  Amer.  Journ.  Sci.,  xvi,  p.  214,  1878. 
Leptoptychaster  arcticus,  var.  elongatus  SLADEN,  op.  cit,  p.  189,  1889. 
Leptoptychaster  arcticus  SLADEN,  op.  cit.,  p.  189.    Verrill,  Proc.  Nat.  Mus., 

xvii,  p.  255,  1894;  Amer.  Journ.  Sci.,  XLIX,  p.  133,  1895.    Ludwig,  Fauna 

Arctica,  i,  p.  452,  1900. 
Leptychaster  arcticus  FISHER,  op.  cit.,  191  ib,  p.  43,  pi.  vm,  fig.  i ;  pi.  ix,  fig.  4. 

This  well-known  Arctic  and  North  Atlantic  species  has  been 
recorded  from  the  North  Pacific  by  Fisher.  It  differs  from  the  other 
west  coast  species  in  having  more  slender  rays  (radial  ratio,  about 
i :  3.25)  and  smaller  supermarginal  plates,  about  forty  on  each  side 
of  a  ray,  which  form  only  a  narrow  margin  to  the  disk.  The  infero- 
marginals  are  quite  oblique  and  have  a  narrow  carina-like  ridge  and 
wide  fasciolate  furrows.  Dr.  Fisher  has  given  a  detailed  description 
of  North  Pacific  specimens. 

In  the  North  Pacific,  according  to  Fisher,  it  ranges  from  Bering 
Sea  to  Yezo,  Japan,  in  72  to  107  fathoms. 

On  the  eastern  coast  of  North  America  it  is  most  frequent  in  80  to 
200  fathoms.  Its  recorded  ranges  in  depth  are  from  50  to  965 
fathoms  (1350  fathoms,  Sladen). 

It  was  taken  by  the  U.  S.  Fish  Commission  parties  at  twenty-three 
stations,  from  N.  lat.  45°  14'  to  38°  29'.  It  always  occurred  in  small 
numbers.  It  is  found  off  the  northern  coasts  of  Europe,  south  to 
Norway  and  off  Ireland,  and  northward  to  the  Arctic  Ocean  and 
Barents  Sea.  It  is  evidently  circumpolar. 

LEPTYCHASTER  PACIFICUS  Fisher. 
Plate  LXXIV,  figure  5. 

Leptychaster  pacificus  FISHER,  Proc.  Wash.  Acad.  Sci.,  vm,  p.  112,  1906;  op. 

cit.,  19116,  p.  45,  pl  vm,  fig.  2;  pi.  ix,  fig.  2;  pi.  L,  figs,  i,  ia. 
Leptychaster  millespina  VERRILL,  op.  cit,  1909,  p.  553. 

Form  regularly  stellate,  with  five  tapering,  acute  rays.  The  radii, 
in  the  specimen  here  described,  are  about  14  mm.  and  35  mm. ; 
ratio,  i :  2.5  ;  often  i :  3. 


SHALLOW-WATER   STARFISHES  327 

It  resembles  L.  arcticus  of  the  North  Atlantic  in  form  and  size,  but 
is  a  more  robust  species,  with  thicker  disk  and  rays  and  larger  and 
fewer  marginal  plates. 

The  marginal  plates  are  about  twenty-two  on  each  side  of  a  ray, 
finely  spinulated,  prominent,  the  lower  ones  projecting  beyond  the 
upper  ones,  and  separated  from  them  by  a  deep  groove;  they 
become  somewhat  oblique  distally.  The  fasciolate  grooves  between 
the  plates  are  deep  and  often  nearly  as  wide  as  the  ventral  plates, 
proximally.  The  lower  marginal  plates  are  high,  compressed,  wedge- 
shaped.  The  dorsal  paxillae  are  small,  close,  and  covered  with 
numerous  fine,  slender  spinules.  When  the  spinules  are  removed 
the  columns  are  small,  unequal,  round,  slender,  with  small  capitate 
tips. 

The  actinal  plates  are  prominent,  subimbricated,  with  deep 
grooves  between ;  their  summits  are  oblique,  elliptical,  covered  with 
numerous  elongated,  slender  spinules. 

The  adambulacral  plates  project  into  the  grooves,  which  are  very 
narrow,  and  have  deep  fasciolate  notches  between  them.  They  bear 
a  furrow-series  of  about  five  or  six  small,  slender,  subequal,  rather 
short  spines,  and  an  actinal  group  of  about  nine  smaller,  divergent 
spinules.  No  pedicellariae  were  found.  Madreporite  is  small  and 
sunken  between  the  paxillae. 

The  two  specimens  here  described  were  taken  at  Departure  Bay, 
British  Columbia,  by  C.  H.  Young,  1908  (Canadian  Geological  Sur- 
vey). Unalaska  to  Straits  of  Georgia  (Fisher),  in  56  to  238 
fathoms. 

Although  this  species  resembles  L.  arcticus  of  the  North  Atlantic, 
it  is  quite  different  in  some  respects.  The  adambulacral  spines  of 
the  latter  are  larger,  fewer,  much  more  unequal ;  the  spinules  of  the 
marginal  plates  and  dorsal  paxillae  are  coarser;  the  marginal  plates 
smaller,  less  prominent  and  more  numerous,  about  forty  to  forty- 
four,  while  in  this,  when  adult,  they  are  from  twenty-two  to  about 
thirty-five.  The  specimen  figured  was  the  type  of  L.  millespina 
Verrill. 

Genus  Glyphaster  Verrill. 

Glyphaster  VESRILL,  Amer.  Naturalist,  xun,  p.  553,   1909.     Fisher,  op.  cit., 
19116,  pp.  39,  53. 

Disk  broad ;  rays  very  short.  Marginal  plates  thick  and  squarish, 
scarcely  oblique.  The  upper  ones  encroach  on  the  upper  surface. 
Each  inferomarginal  corresponds  to  two  adambulacrals  on  the  rays, 
and  nearly  three  in  the  interradial  areas.  The  marginal  plates  are 


328  VERRILL 

covered  with  fine  spinules.  The  inferomarginal  plates  are  convex, 
not  carinated.  The  dorsal  plates  are  finely  spinulated  paxillae. 
Adambulacral  plates  much  as  in  Leptychaster. 

Professor  Fisher  (op.  cit,  191  ib)  does  not  recognize  this  as  a 
distinct  genus. 

GLYPHASTER  ANOMALUS  (Fisher)  Verrill. 
Plate  i,  figures  i,  2;  plate  vi,  figures  i,  2. 

Leptychaster  anotnalus  FISHER,  1906,  p.  115;  ignb,  p.  48,  pi.  vn,  fig.  4;  pi.  ix, 

fig.  i;  pi.  L,  figs.  2,  20. 
Glyphaster  anotnalus  VERRILL,  op.  cit,  1909,  XLIH,  p.  554. 

The  stout  inferomarginal  plates  on  the  disk  and  rays  correspond 
each  to  two  adambulacrals,  except  the  first  pair,  each  of  which  is 
connected  with  three  adambulacrals  by  lines  of  interactinal  plates. 
But  in  L.  arcticus  each  inferomarginal  corresponds  pretty  closely 
with  a  single  adambulacral,  except  the  two  proximal  interradials, 
each  of  which  is  connected  with  two  adambulacrais  by  lines  of  inter- 
radials. 

The  sutures  between  the  inferomarginals  are  straight  and  distally 
nearly  at  right  angles  to  the  furrows,  while  in  L.  arcticus  they  are 
all  oblique,  and  distally  very  oblique  and  curved,  with  the  inner  end 
turned  adorally,  so  that  the  corresponding  adambulacral  is  about 
opposite  the  outer  end  of  the  preceding  marginal. 

The  epioral  side  of  the  jaws  is  elevated  and  laterally  compressed, 
narrower  aborally,  and  the  two  halves  are  close  together,  leaving  a 
narrow,  strongly  fasciolated  groove  between  them,  while  in  L.  arcti- 
cus they  are  not  so  prominent,  elliptical,  with  no  distinct  fasciolated 
groove. 

Bering  Sea,  off  Pribilof  Islands  and  St.  Paul,  to  southern  Alaska, 
and  to  Japan,  in  32  to  688  fathoms;  off  Monterey,  California,  in 
871  fathoms  (Fisher). 

Family  PORCELLANASTERID2E  Sladen. 

Porcellanasteridce  SLADEN,  1886 ;  Voy.  Chall.,  xxx,  p.  125,  1889.    Fisher,  op.  cit., 
19116,  p.  22. 

Disk  usually  broad,  with  short  rays  and  high  margin.  Marginal 
plates  usually  paired  (inferior  ones  are  sometimes  abortive),  thin, 
usually  covered  with  strong  membrane,  seldom  granulose,  sometimes 
with  spines;  edges  commonly  fasciolated,  or  there  may  be  special 
fasciolated  or  cribriform  organs. 


SHALLOW-WATER    STARFISHES  329 

Dorsal  ossicles  either  paxilliform,  or  simple  plates  with  a  single 
spinule.  Interactinal  plates  flattened,  imbricated.  Adambulacral 
armature  simple,  uniserial.  Ampullae  single.  No  intestine.  Dorsal 
pore  generally  absent.  Superambulacral  plates  usually  absent. 

The  stomach  is  very  large  ;  the  food  consists  of  fine  mud  or  sand 
containing  foraminifera,  etc.,  which  these  starfishes  swallow  in  large 
quantities.  Hence  they  are  best  adapted  to  life  on  bottoms  of  "  Glo- 
bigerina  ooze,"  in  the  deep  seas. 

This  family  was  divided  by  Sladen,  1886,  into  two  subfamilies: 
Porcellasterinae  and  Ctenodiscinae. 

PORCELLASTERIN,E  Sladen. 

In  these  there  are  special  localized  cribriform  organs.  No  fascio- 
lated  channels  between  the  interactinal  plates.  Superambulacrals 
always  or  nearly  always  lacking.  All  are  confined  to  deep  water. 


CTENODISCIN;E 

In  these  there  are  simple  fasciolated  channels  between  the  inter- 
actinal plates  and  continuous  with  more  definite  ones  between  the 
marginals,  covered  by  small,  flat  spinules.  (See  pi.  v,  figs.  6-80.) 
Dorsal  plates  are  paxilliform.  Superambulacrals  are  present 

Subfamily  CTENODISCIN&  Sladen. 

Ctenodiscina  (subfamily  of  Porcellanasteridce)   SLADEN,  op.  cit.,  1886;  Voy. 
Chall.,  xxx,  pp.  127,  170,  1889. 

Disk  broad,  flat,  with  short  rays.  Marginal  plates  thin,  covered 
with  membrane,  not  granulose,  paired  and  conterminal,  in  two 
closely  united  rows,  furnished  laterally  with  specialized  fasciolate 
grooves,  covered  by  a  single  row  of  flattened  spinelets.  Disk  usually 
with  true  paxillae;  paxillar  columns  enlarged  at  base,  but  isolated. 
Actinal  interradial  areas  covered  with  flat  plates,  with  fasciolated 
grooves  between  the  rows  or  double  rows,  continuous  with  those 
between  the  marginal  plates  and  adambulacral  plates. 

An  unpaired  interradial  marginal  plate,  in  each  series,  is  present 
in  Pectinodiscus  Ludwig  ;  also  an  odd  row  of  actinal  plates. 

Adambulacral  plates  with  a  divergent  group  of  furrow-spines,  and 
some  on  the  actinal  side.  A  minute  central  dorsal  pore,  often  on  an 
elevation. 

At  present  this  subfamily  includes  only  Ctenodiscus  and  Pectino- 
discus Ludwig.  It  is  nearly  intermediate  in  many  respects  between 


33O  VERRILL 

Porcellanasterinas  and  some  of  the  Astropectinidae,  especially  Lepty- 
chaster.  From  the  latter  it  differs  in  the  somewhat  more  specialized 
fascicles  of  the  marginal  and  actinal  plates,  and  their  thinner  struct- 
ure and  naked  central  portions.  From  the  Porcellanasterinae  the  dif- 
ferences seem  important.  The  latter  have  much  more  specialized 
fascicles,  in  special  locations  on  the  margin,  with  none  between  the 
actinal  plates,  and  usually  lack  regular  dorsal  paxillae.  Like  the 
latter,  it  has  single  ambulacral  ampullae. 

Genus  Ctenodiscus  Muller  and  Troschel. 

Ctenodiscus  MULLER  and  TROSCHEL,  Syst  Ast,  p.  76,  1842.     Sladen,  op.  cit, 
p.  170,  1889.    Fisher,  op.  cit.,  jgnb,  p.  31. 

Stellate  with  nearly  vertical  margins.  Actinal  plates  thin  and  flat, 
without  granules ;  covered  with  membrane ;  the  principal  ones  united 
in  double  radial  rows,  each  double  row  corresponding  to  a  single 
marginal  plate  and  to  two  adambulacral  plates.  Thus  the  fasciolate 
grooves  run  between  the  double  rows  (or  alternate  single  rows). 
But  distally,  on  the  rays,  the  adambulacral  and  marginal  plates  cor- 
respond, plate  for  plate. 

Dorsal  ossicles  are  true  paxillae.  The  upper  marginal  plates  are 
high,  thin,  nearly  vertical,  and  proximally  form  a  narrow  margin, 
usually  with  a  small  spine  to  each  plate;  but  distally  they  encroach 
upon  the  upper  side  of  the  rays. 

The  central  dorsal  area  often  has  a  more  or  less  conical  elevation 
of  the  dermis,  covered  with  paxillae  like  the  adjacent  parts,  but  finer. 
It  is  more  prominent  in  the  young,  and  when  the  stomach  is  filled 
with  soft  mud,  as  is  usually  the  case  when  living. 

The  northern  and  arctic  species  (C.  crispatus)  is  circumpolar. 
Two  closely  allied  species  or  varieties  are  found  in  Antarctic  waters, 
one  on  each  coast  of  southern  South  America.  Dr.  Fisher  thinks 
that  they  may  not  be  distinct  from  C.  crispatus. 

CTENODISCUS  CRISPATUS  (Retz.)  Diiben  and  Koren. 
Plate  v,  figures  6,  7,  8,  8a  (details)  ;  plate  XLIX,  figures  5,  50  (details). 

Asterias  crispatus  RETZIUS,  Dissert.  Asteriarum,  p.  17,  1805. 

Ctenodiscus  polaris  and  C.  pygmaus  MULLER  and  TROSCHEL,  Syst,  pp.  76,  129, 

pi.  v,  fig.  5,  1842. 
Ctenodiscus  crispatus  DUBEN  and  KOREN,  K.  Vet.  Akad.  Handl.,  p.  253,  1844. 

Stimpson,  Invert.  G.  Manan,  p.  15,  1853.    Ltitken,  Gronl.  Echinod.,  p.  45, 

1857.     Verrill,    Proc.   Bost.    Soc.   Nat.   Hist,   x,   p.   345,    1866;    Amer. 

Naturalist,  XLIII,  p.  548,  549,  figs,  3,  a,  b  (4-rayed  and  5-rayed),  1909. 


SHALLOW- WATER   STARFISHES  331 

Ctenodiscus  corniculatus  PERKIER,  Stell.  du  Mus.,  p.  380,  1875  (after  Linck). 

Duncan  and  Sladen,  Echinod.  Arctic  Sea,  p.  49,  pi.  in,  figs.  17-20,  1881. 

Sladen,  Voy.  Chall.,  p.  171,  1889.     Doderlein,  op.  cit,  p.  221,  pi.  ix,  figs. 

2,  3,  1900. 
Ctenodiscus  krausei  LUDWIG,  Zool.  Jahrb.,  Syst.,  i,  p.  290,  pi.  vi,  figs.  13-16, 

1886. 
Ctenodiscus  crispatus  FISHER,  op.  cit.,  I9II&,  p.  31,  pi.  in,  figs.  1-4;  pi.  iv,  figs. 

1-6. 

The  following  description  is  from  Alaskan  specimens.  Disk 
broad,  thin,  with  regularly  curved  interradial  margins,  which  are 
bordered  by  the  summits  of  the  thin  upper  ends  of  the  upright 
superomarginal  plates,  each  of  which  usually  bears  a  small  conical 
spine. 

Rays  five  (rarely  four),  broad  at  base,  rapidly  tapered,  subacute, 
varying  considerably  in  length,  relative  to  disk.  Radii  of  a  specimen 
from  off  Chilikoff,  Alaska,  are  14  mm.  and  24  mm. ;  ratio,  i :  1.7. 

Another,  from  the  same  place,  has  the  radii  12  mm.  and  23  mm. ; 
ratio,  i:  1.9.  These  are  of  medium  proportions;  the  ratios  of  the 
radii  are  often  i :  2,  and  up  to  i :  2.25. 

The  dorsal  paxillae  are  minute  and  pretty  uniformly  crowded  over 
the  whole  surface.  Madreporite  rather  large  and  conspicuous. 
Ocular  plate  rather  large,  shield-shaped,  longer  than  broad,  distal 
end  notched. 

The  upper  marginal  plates  are  closely  united  to  the  lower  ones, 
end  to  end,  so  that  the  fascioles  between  them  coincide;  those  in  the 
interradial  areas  extend  but  slightly  on  the  disk,  but  the  small  distal 
ones  encroach  considerably  on  the  upper  side  of  the  rays.  The  mar- 
ginal spine  is  short,  subacute,  articulated  on  a  central  mammilliform 
tubercle,  which  shows  a  central  pit  when  the  spine  is  removed.  The 
tubercle  is  wider  than  the  median  naked  keel  of  the  plate,  except 
distally  on  the  rays,  where  the  keel  becomes  wider.  The  fasciolated 
spinules  are  longer  than  the  width  of  the  keel,  except  distally  on  the 
rays ;  they  are  flat,  and  project  at  nearly  right  angles  to  the  edges  of 
the  keel  and  parallel  to  the  surface  of  the  plate ;  they  form  a  single 
regular  close  row,  with  a  small,  well  defined  channel  under  them. 
This  channel  extends  down  between  the  inferomarginal  plates,  with 
the  same  sort  of  fasciolated  spinules ;  but  the  latter  gradually  become 
shorter  downward,  and  point  obliquely  upward. 

The  inferomarginal  plates  of  the  interradial  arcs  are  considerably 
shorter,  vertically,  than  the  upper  ones,  and  have  the  naked  median 
keel  more  than  twice  as  wide,  while  toward  the  ends  of  the  rays  it 
becomes  much  wider,  covering  most  of  the  squarish  plates,  the 


332  VERRILL 

fasciolated  spinules  becoming  very  short.  These  distal  plates  have  a 
slightly  granular  surface,  but  no  articulated  granules.  Most  of  the 
inferomarginals  bear  a  small,  subacute,  articulated  spine  or  tubercle 
near  the  upper  end. 

The  actinal  plates,  in  specimens  of  ordinary  adult  size,  form  six 
double  rows,1  running  from  the  six  interradial  inferomarginal  plates 
to  twelve  adambulacral  plates.  The  plates  are  small  next  the  mar- 
ginals, but  become  much  larger  toward  the  adambulacrals ;  they  are 
smooth,  flat,  irregularly  angular,  with  rounded  corners,  and  alternate 
closely  in  the  two  rows,  without  intervening  grooves  or  spinules; 
their  united  edges  overlap  slightly. 

Sometimes  there  are  slight  irregularities  in  this  arrangement;  the 
more  distal  of  the  double  rows  may  correspond  with  but  one  adambu- 
lacral, on  some  of  the  rays.  Farther  out  the  marginal  and  adambu- 
lacral plates  correspond,  and  there  are  but  two  (or  only  one)  actinal 
plates  between  them.  Beyond  the  sixth  marginal,  the  remaining 
five  or  six  are  joined  directly  to  the  adambulacrals. 

The  fasciolated  grooves  run  directly  from  between  all  the  infero- 
marginal plates  to  the  ambulacral  grooves,  thus  passing  between  the 
double  rows  of  actinals ;  and  distally  between  all  the  marginals  and 
adambulacral  plates.  They  are  bordered  by  regular,  close,  single 
rows  of  short  appressed  spinules. 

The  adambulacral  plates  are  large  and  have  a  strong  triangular 
lobe  projecting  into  the  grooves,  ending  in  line  with  a  sharp  ridge 
that  stands  between  the  large  ambulacral  feet.  The  proximal  plates 
bear  a  series  of  three  or  four  (sometimes  five)  acute,  divergent 
furrow-spines;  the  distal  ones,  only  two  spines;  on  the  actinal  side 
there  may  be  three  to  five  small  spines,  mostly  marginal.  Oral  spines 
numerous,  the  two  medial  adorals  stouter. 

Color,  light  yellow  to  pale  orange,  the  upper  surface  often  gray 
when  the  stomach  is  distended  with  mud,  as  it  frequently  is. 

The  median  dorsal  nephridial  cone  is  usually  low  in  the  adults, 
but  elongated  in  the  very  young  ones.  The  central  pore  is  minute. 

I  have  studied  a  considerable  number  of  regular  four-rayed 
specimens  taken  off  the  coast  of  New  England  by  myself  and  by  the 
U.  S.  Fish  Commission. 

This  species  is  abundant  in  the  North  Atlantic  and  Arctic  Oceans, 
on  soft  muddy  bottoms,  in  from  10  to  650  fathoms.  It  is  found  on 

1The  detailed  figure  of  these  plates,  given  by  Miiller  and  Troschel,  is  in- 
correct, as  to  their  regular  arrangement  in  double  and  single  columns.  (See 
pi.  XLIX,  fig.  5,  copied  from  Mtiller  and  Troschel,  and  pi.  v,  fig.  6.) 


SHALLOW-WATER   STARFISHES  333 

the  east  American  coast  from  Greenland  south  to  Cape  Cod  in 
shallow  water,  and  in  deep  water  to  the  latitude  of  Cape  Hatteras. 
Particularly  abundant  in  the  Gulf  of  Maine  and  Massachusetts  Bay, 
in  20  to  100  fathoms,  where  several  hundreds  are  often  taken  at  one 
haul  of  the  dredge.  It  is  also  common  on  the  northern  coasts  of 
Europe,  south  to  Norway  and  the  Faroe  Islands.  I  have  seen  a 
number  of  specimens  from  northern  Alaska.  Off  Chilikoff  (Pro- 
fessor Kincaid),  described  above,  and  from  Vancouver  Island. 

C.  krausei  Ludwig  was  described  from  two  small  specimens  from 
Bering  Sea.  Both  specimens  had  the  larger  radius  18  mm;  lesser, 
9.5  mm.  Subsequently  Ludwig  himself  referred  them  to  C.  crispa- 
tus.  According  to  his  figure  (see  pi.  v,  fig.  6),  they  differ  from 
those  described  above  in  having  only  two  double  columns  of  actinal 
plates  in  the  interradial  areas,  due,  perhaps,  to  immaturity. 

Fisher  records  it  from  numerous  localities  in  the  North  Pacific, 
mostly  from  Bering  Sea  and  off  Alaska,  southward  to  California,  in 
31  to  1033  fathoms,  and  off  the  Asiatic  coast  to  Japan.  Off  Gulf  of 
California  and  Panama  (Ludwig).  Dr.  Fisher  also  believes  that  the 
forms  described  from  off  the  two  coasts  of  southern  South  America 
as  C.  australis  and  C.  procurator,  by  Sladen,  are  not  distinct.  If  this 
be  true,  its  range  would  be  greatly  extended. 

It  is  a  very  variable  species,  if  all  the  forms  belong  to  one  species. 

Family  LUIDUD2E  Verrill. 

f.uidiida1  VERRILL,  Trans.  Conn.  Acad.,  x,  p.  201,  1899.    Fisher,  op.  cit,  191 1&, 

p.  105. 
Luidiina  (subfamily)  SLADEN,  op.  cit.,  pp.  175,  244,  1889. 

Disk  small,  covered  with  true  paxillae,  which  are  usually  largest  at 
the  sides  of  disk  and  rays ;  rays  five  to  ten  or  more,  long,  flat,  flexible 
in  life ;  upper  marginal  plates  small,  paxillif orm  or  indistinguishable ; 
lower  ones  large,  spinose,  fascicled,  corresponding  in  number  and 
length  with  the  adambulacrals  and  separated  from  them  throughout 
the  rays  by  a  small  peractinal  plate. 

The  dorsal  nephridial  glands,  or  "  coecal  appendages,"  intestine 
and  dorsal  pore  are  lacking;  the  superomarginal  plates  are  small, 
usually  not  easily  distinguishable.  The  papulae  are  branched. 

The  gonads  are  multiple  and  arranged  in  rows,  with  separate  pores 
along  the  sides  of  the  rays. 

Pedicellariae  usually  present  on  the  actinal  side,  usually  forceps- 
like,  with  two  or  three  blades,  or  spiniform. 


334  VERRILL 

Ambulacral  feet  large,  flattened,  muscular ;  can  be  used  as  paddles 
for  gliding  rapidly  along  the  bottom  or  just  under  the  surface  of 
sand  or  mud. 

This  family  seems  eminently  worthy  of  separation  from  Astropec- 
tinidae.  Its  whole  structure  is  adapted  to  its  life  under  the  surface 
of  sand  or  mud,  and  for  rapid  motion.  I  have  observed  that  Luidia 
clathrata  (see  pi.  cm,  fig.  i)  swims  or  paddles  with  remarkable  speed, 
just  under  the  surface  of  the  sand  in  shallow  water,  and  that  it  swims 
or  glides  actively  in  an  aquarium,  by  using  its  feet  as  paddles.  ( See 
P.  7-) 

Genus  Luidia  Forbes. 

Luidia  FORBES,  Wern.  Trans.,  1839,  p.  14 ;  Mem.  Wern.  Soc.,  vm,  p.  128,  1840. 
Muller  and  Troschel,  Syst.  Aster.,  p.  77,  1842.  Sladen,  op.  cit,  p.  244, 
1889,  Fisher,  191  ib,  p.  105. 

Rays  five  to  ten,  rather  flat,  flexible.  Dorsal  columnar  paxillae 
have  lobate,  articulated  bases;  summits  either  plainly  paxillose  or 
with  a  central  spine  or  tubercle  (spinopaxillae),  or  with  both  kinds; 
largest  next  the  lateral  borders  of  the  rays.  Inferomarginal  plates 
spinose  and  spinulose.  Pedicellariae  often  absent;  when  present, 
forceps-like,  with  two  or  three  blades  or  valves.  They  are  usually 
situated  on  the  adambulacral  plates;  sometimes  on  the  inferomar- 
ginals  or  peractinals. 

Inferomarginal  plates  transversely  elongated,  with  wide  and  deep 
fasciolated  grooves  between  them,  bordered  by  slender  spinules; 
central  portion  with  one  or  more  rows  of  spines.  Superomarginals 
small,  paxilliform,  rounded.  Adambulacral  plates  short,  with  one 
furrow-spine  (rarely  two)  ;  they  are  separated  by  wide  grooves. 
The  dorsal  paxillae  may  be  quadrate  at  surface  and  crowded,  in 
regular  rows,  or  stellate.  Regular  rows  of  papulae  between  the  rows 
of  paxillae.  No  superambulacral  plates  observed. 

LUIDIA  FOLIOLATA  Grube. 
Plate  c,  figures  2,  20;  plate  cm,  figure  2;  plate  cv,  figure  2. 

Luidia  fotiolata  GRUBE,  op.  cit.,  Breslau,  XLIII,  p.  69,  1866.    Fischer,  op.  cit., 
19116,  p.  106,  pi.  xix,  figs.  1-3;  pi.  xxi,  figs.  3-5;  pi.  LIV,  fig.  3. 

This  species  grows  to  large  size  on  the  California  coast.  One 
from  British  Columbia  has  the  radii  25  mm.  and  205  mm.;  ratio, 
i :  8.2.  Another  has  the  radii  18  mm.  and  130  mm. ;  ratio,  1 :  5.5. 
The  rays  are  rather  flat  proximally,  gradually  tapered  to  slender  tips. 


SHALLOW-WATER   STARFISHES  335 

The  paxillae  form  regular  longitudinal  and  transverse  rows. 
Those  next  the  margin  are  rather  quadrate  superficially ;  the  central 
ones  are  small,  crowded,  roundish,  stellate;  all  have  flat  or  slightly 
convex  summits.  The  larger  are  i  mm.  to  1.25  mm.  broad;  they 
have  a  central  rosette  of  small  clavate  or  capitate  spinules,  of  which 
one,  or  sometimes  two,  stand  at  the  center  and  six  or  eight  around 
it ;  outside  of  this  is  a  marginal  horizontal  fringe  of  longer  and  more 
slender  spinules,  twelve  to  fifteen  or  more  in  number,  making 
fasciolated  spaces.  When  the  spinules  are  removed,  the  paxillary 
column  has  a  somewhat  enlarged,  roundish  or  oblong-elliptical  sum- 
mit; the  base  is  regularly  four-lobed;  the  lobes  are  carinate  and 
articulated  movably.  Between  the  rows,  alternating  with  the  paxillae, 
are  regular  rows  of  large  papular  pores.  Proximally  there  are,  on 
each  side,  about  five  or  six  rows  of  paxillae. 

The  upper  marginal  plates  are  paxilliform  and  similar  to  the 
adjacent  paxillae,  but  closely  joined  to  the  upper  end  of  the  infero- 
marginals;  distally  they  become  relatively  larger. 

The  inferomarginal  plates  are  large  and  prominent,  their  upper 
ends  defining  the  margin.  Each  bears  about  three  to  five  principal 
spines,  the  outer  ones  larger,  crowded,  flattened,  obtuse;  the  lower 
ones  smaller  and  more  acute.  Besides  these  there  are  numerous 
slender,  sharp,  unequal  spines  below,  and  many  marginal  spinules. 

The  peractinal  row  of  ossicles  is  continuous  with  the  inferomar- 
ginals,  and  each  bears  a  cluster  of  slender  spinules,  like  those  of  the 
latter. 

The  adambulacral  plates  have  an  inner,  bent,  angular  or  triquetral, 
subacute  spine  and  about  three  slightly  smaller,  slender,  roundish, 
tapered,  straight  spines  in  a  triangular  group  on  the  actinal  surface. 

A  few  small  actinal  plates,  near  the  mouth,  bear  small,  slender 
spines.  A  few  slender  spinuliform  pedicellariae,  of  diverse  sizes, 
were  noticed  on  the  adambulacral  plates  and  on  the  interradial  areas. 

Color,  mottled  gray  above,  pale  yellow  below. 

This  species  is  not  uncommon  on  the  coast,  in  shallow  water,  from 
San  Francisco,  California,  to  British  Columbia.  Departure  Bay,  in 
17  to  25  fathoms,  gravel  and  sand  (C.  H.  Young,  1908,  Canadian 
Geological  Survey)  ;  Pacific  Grove,  California  (Prof.  W.  R.  Coe)  ; 
off  San  Francisco  (Prof.  W.  E.  Ritter)  ;  Victoria,  Vancouver 
Island  (C.  F.  Newcombe). 

Dr.  Fisher  gives  its  distribution  as  from  Southern  Alaska  to  San 
Diego,  California,  in  10  to  189  fathoms,  mostly  less  than  80  fathoms. 
He  had  it  from  a  large  number  of  localities. 


336  VERRILL 

LUIDIA  LUDWIGI  Fisher. 

Luidia  ludwigi  FISHER,  op.  cit.,  1906,  p.  122;  op.  cit.,  191  ib,  p.  113,  pL  xx, 
figs.  2,  3;  pi.  xxi,  fig.  2;  pi.  LIV,  figs.  2,  20. 

Rays  five,  long  and  narrow.  Radii,  13  mm.  and  107  mm. ;  ratio, 
i :  8.2.  Abactinal  area  of  rays  has  three  or  four  rows  of  quadrate 
paxillae  on  each  side. 

The  superomarginal  plates  have  small  trivalve  and  bivalve  pedi- 
cellariae. 

The  inferomarginals  have  a  similar  pedicellaria  on  the  upper  end, 
and  one  to  three,  mostly  two,  lateral  spines,  and  three  to  six 
enlarged  spinelets,  on  the  lower  side,  with  small  spinelets  over  the 
general  surface. 

The  peractinal  plates  extend  to  half  the  length  of  the  ray;  and 
like  those  of  the  interradial  areas,  each  has  a  rather  prominent  three- 
valved  pedicellaria. 

The  adambulacral  plates  have  a  curved  furrow-spine,  three  spines 
on  the  actinal  side,  and  one  or  two  smaller  spines  ( Fisher) . 

According  to  Dr.  Fisher,  this  species  occurs  from  Monterey  Bay 
to  Santa  Barbara  Islands  and  San  Pedro,  California,  in  15  to  50 
fathoms,  mostly  on  bottoms  of  mud  or  sand. 

LUIDIA  ASTHENOSOMA  Fisher. 

Luidia  asthenosoma  FISHER,  op.  cit.,  1906,  p.  124;  IQII&,  p.  116,  pi.  xx,  fig.  i;  pi. 
xxi,  fig.  i ;  pi.  LIV,  fig.  i. 

Rays  five,  very  long,  slender,  flattened,  fragile,  with  a  small  disk. 
Radii,  9  mm.  and  86  mm. ;  ratio,  i :  9.5 ;  width  of  ray  at  base. 
10  mm.  to  ii  mm. 

Inferomarginal  plates  narrow,  forming  the  margins  of  the  ray, 
but  not  extending  much  on  the  dorsal  surface,  each  with  a  transverse 
row  of  three  large,  sharp,  acicular  spines,  and  covered  with  very 
slender  spinules ;  fascicles  deep  and  wide. 

The  adambulacral  plates  have  a  transverse  row  of  three  prominent 
spines,  the  inner  one  being  a  saber-shaped  furrow-spine.  The  inter- 
actinal  or  peractinal  plates  have  a  bivalved,  short,  blunt,  papilliform 
pedicellaria. 

The  dorsal  paxillae  are  stellate;  some  have  two-valved,  or  some- 
times three-valved,  pedicellariae.  The  superomarginal  plates  are 
about  twice  the  size  of  the  adjacent  paxillae.  The  oral  plates  have 
bivalved  pedicellariae  (Fisher). 

Dr.  Fisher  records  this  species  from  thirty- four  localities,  in  1 1  to 
339  fathoms,  between  Monterey  Bay  and  Los  Coronados  Islands, 
Lower  California. 


SHALLOW-WATER   STARFISHES  337 

SPECIES  WRONGLY  ATTRIBUTED  TO  THIS  FAUNA. 

Asterias  rubens  Linn. — Ives  (1890)  recorded  this  from  San  Luis 
Obispo.  Murdoch  recorded  it  from  Point  Barrow,  Arctic  Ocean. 
The  latter  proves  to  be  Allasterias  rathbuni  Verrill. 

Asterias  lurida  Phil. — Ives  (1889)  recorded  this  from  San  Diego, 
California.  It  is  a  Chilean  species  and  not  likely  to  be  found  in 
Californian  waters.  The  species  mistaken  for  it  is  uncertain. 

.Linckia  guildingii  Gray. — This  species  is  known  only  from  the 
West  Indies  and  Bermuda.  It  was  recorded  with  doubt  by  Ives 
( 1889)  from  San  Diego.  Probably  his  small  specimen  was  the  young 
of  L.  Columbia. 

For  southern  (Panamic}  species  recorded  from  San  Diego  or  fur- 
ther north,  see  below,  p.  346.  Perhaps  some  are  errors  as  to  locality 
due  to  misplaced  labels. 

GEOGRAPHICAL  DISTRIBUTION. 

The  following  account  of  the  geograhpical  distribution  of  the 
shallow-water  species  included  in  this  paper  is  intended  merely  to 
give  the  general  facts,  so  far  as  now  known  to  me.  No  doubt  the 
elaboration  of  the  extensive  collections  of  Forcipulata  made  by  the 
steamer  Albatross,  from  deeper  waters,  will  very  materially  change 
the  range  of  many  species  now  known  only  from  the  shores  and 
very  shallow  waters. 

For  my  present  purpose,  the  entire  region  may  be  divided  into 
four  great  faunal  districts: 

I.  The  Beringian  or  North  Alaskan. — This  includes  the  Arctic 
Ocean  coast  of  Alaska  and  all  the  coasts  and  islands  of  Bering  Sea, 
south  to  the  Aleutian  Islands  and  Alaskan  Peninsula. 

II.  The  Columbia-Alaskan. — This  includes  the  coasts  and  islands 
of  Alaska,  south  of  the  Aleutian  Islands,  and  the  entire  coast  of 
British  Columbia,  with  Vancouver  Island;  Puget  Sound,  and  the 
northwestern  coast  of  Washington;  the  Gulf  of  Georgia;  and  the 
Straits  of  Fuca. 

III.  The  Californian. — This  includes  the  middle  and  southern 
parts  of  the  coast  of  Washington ;  all  of  the  Oregon  coast ;  and  the 
coast  of  California  to  Point  Conception,  or  the  north  end  of  the 
Santa  Barbara  Channel. 

IV.  The  South  Californian. — This  includes  the  coast  of  southern 
California,  from  the  Santa  Barbara  Channel,  Santa  Rosa  Island, 
and  Santa  Cruz  Island  to  the  middle  part  of  the  Lower  Californian 
coast. 

23 


VERRILL 

This  fauna  was  not  originally  considered  as  pertaining  to  this 
report,  but  some  of  its  species  were  introduced  for  special  reasons. 
The  list,  however,  is  not  thought  to  be  by  any  means  complete. 

I.  THE  BERINGIAN  FAUNA. 

The  following  species  have  occurred  in  less  than  60  fathoms. 
Those  with  an  asterisk  prefixed  occur  also  in  the  North  Atlantic,  and 
are  believed  to  be  circumpolar. 

Those  with  a  dagger  prefixed  belong  more  properly  to  the 
Columbia-Alaskan  fauna,  and  have  their  northern  limits  in  the 
southern  part  of  Bering  Sea. 

*Asterias  acervata  St. 

Arctic  Ocean;  Bering  Sea;  Bering  Strait. 
Asterias  multiclava  Ver. 

Arctic  Ocean ;  Bering  Sea ;  Bering  I. ;  Siberia. 
A.  polythela  Ver. 
Arctic  Ocean. 
Leptasterias  obtecta  Ver. 
Bering  Sea,  17  fath. 
L.  arctica  (Murdoch). 

Arctic  Ocean;  Point  Barrow. 
fL.  epichlora  alaskensis  Ver. 

Aleutian  Islands  southward  to  Vancouver  I. 
*Ctenasterias  cribraria  (St.)  Ver. 

Arctic  coast  of  Alaska. 
*Stephanasterias  albula  (St.)  Ver. 

Bering  Sea ;  Arctic  Ocean. 
Allasterias  rathbuni  Ver. 

Unalaska;  Bering  Sea. 
A.  rathbuni  nortonensis  Ver. 

Norton  Sd. 

Allasterias  anomala  Ver. 
Off  St.  Michael's  I. 
•\Henricia  leviuscula  (St.). 

Bering  I.  and  Aleutian  Is.  south  to  southern  California. 
•\H.  leviuscula,  var.  multispina  F. 

Bering  I.,  southward  to  Puget  Sound,  0-238  fathoms. 
fH.  leviuscula,  var.  spiculifera  (Clark). 

Aleutian  Is.  to  Puget  Sound. 
*H.  sanguinolenta  (Mull.). 

Arctic  coast  south  to  the  Aleutian  Is. ;  Point  Barrow ;  off  Washington 

(Fisher). 
*Var.  pectinata  Ver. 

Arctic  Ocean  and  Bering  Sea;  41-349  fathoms. 
Var.  rudis  Ver. 

Point  Franklin,  Arctic  Ocean,  13}^  fathoms. 
H.  tumida  Ver. 

Arctic  Ocean  to  Dutch  Harbor,  Alaska. 


SHALLOW-WATER   STARFISHES  339 

H.  tumida  borealis  Ver. 

Bering  Str.,  south  to  Yakutat  and  Sitka,  Alaska ;  Siberia. 
H.  arctica  Ver. 

Cape  Lisburne. 
fH.  aspera  F. 

Bering  Sea  to  Santa  Barbara,  Calif. 
*Solaster  endeca  (L.). 

Arctic  Ocean ;  Bering  Sea,  south  to  Sitka ;  Queen  Charlotte  Sd.,  238  fath. 
fS.  dawsoni  Ver. 

Kuril  Is. ;  Aleutian  Is.,  southward  to  California. 
5".  dawsoni  arctica  Ver. 

Point  Franklin,  Arctic  Ocean. 
f5".  stimpsoni  Ver. 

Bering  I.  and  Commander  Is.,  south  to  Oregon. 
S.  paxillatus  Sladen. 

Bering  Sea  to  Kadiak  and  to  Japan;  56-350  fath. 
*Crossaster  papposus  (L.). 

Cape  Franklin,  Arctic  Ocean;  Bering  Sea  to  Oregon;  Siberia;  common 

off  Alaska,  1-283  fath. 
^Pteraster  tesselatus  Ives. 

Bering  I.  to  Puget  Sd. 
*P.  wilitaris  Mull. 

Bering  Str.  to  Aleutian  Is. ;  Str.  of  Fuca,  100  fath. 
*P.  pulvillus  Sars. 

Off  Bering  I.,  in  deep  water. 
P.  marsippus  F. 

Bering  Str.,  52-351  fath. 
*Pter aster  obscurus  (Per.). 

Bering  Str.  to  Unalaska  and  Kamchatka. 
P.  obscurus,  var.  octaster  Ver. 

Bering  Str.  to  Pribilof  Is.  and  Kamchatka. 
Pterasterides  aporus  (Lud.). 

Bering  Sea. 
*Diplopteraster  multipes  (Sars)  Ver. 

Bering  Sea  to  Aleutian  Is.,  81-350  fath.;  off  San  Diego,  Calif.,  628-640 

fath. 
*Ceramaster  granularis  (R.)  Ver. 

Bering  Sea  to  Aleutian  Is.;  Str.  of  Georgia  (?). 
C.  patagonicus  (Sla.)  F. 

Bering   Sea  to   Aleutian   Is.   and  Unalaska;    Southern   Alaska,   41-134 

fath.;  Gulf  of  California  (Fisher). 
Tosiaster  arcticus  Ver. 

Bering  Sea  to  Aleutian  Is.  and  Kadiak,  Alaska,  and  to  Siberia;  0-102 

fath. 
^Hippasteria  spinosa  Ver. 

Bering  Sea  to  southern  California. 
Luidiaster  dawsoni  (Ver.). 

Bering  Sea  to  British  Columbia,  56-159  fath. 
*Leptychaster  arcticus  (Sars). 

Bering  Sea  to  Bering  I.,  and  to  Yezo,  Japan 


34°  VERRILL 

L.  pacificus  F. 

Bering  Sea  to  Vancouver  I.  and  to  Str.  of  Georgia. 
Glyphaster  anomalus  (Fisher)  V. 

Bering  Sea  to  southern  Alaska  and  Japan ;  32-688  fath. 
*Ctenodiscus  crispatus  (Retz.)  M.  &  Tr. 

Bering  Sea  to  California  and  Japan;  31-1033  fath. 

II.  THE  COLUMBIA-ALASKAN   FAUNA. 

This  extensive  region  comprises  by  far  the  longest  list  of  species. 
Those  with  an  asterisk  prefixed  belong  more  properly  to  the  Bering- 
ian  fauna  but  extend  southward,  often  far  into  this,  in  deep  water. 
Those  with  a  dagger  (f)  prefixed  are  more  southern  species  that 
find  their  northern  limits  in  the  southern  part  of  this  fauna.  Many 
species  are  common  to  this  and  the  next  fauna. 

Pisaster  ochraceus  (Br.)  Ag. 

Aleutian  Is.  and  Yakutat  to  Monterey  Bay. 
Var.  nodiferus  Ver. 

Sitka;  British  Columbia  to  Gulf  of  Georgia. 
P.  confer tus  (St.)  Ver. 

British  Columbia  to  Puget  Sd.  and  Vancouver  I. 
P.  liitkenii  (St.)  Ver. 

Vancouver  I.  to  Monterey  Bay;  and  as  var.  australis,  to  San  Diego. 
P.  paucispinus  (St.)  Ver. 

British  Columbia  and  Puget  Sd.  to  California. 
P.  papulosus  Ver. 

Departure  Bay,  British  Columbia,  and  Puget  Sound  to  Gulf  of  Georgia. 
P.  brevispinus  (St.)  Ver. 

Washington  to  Monterey,  California. 
Asterias  victoriana  Ver. 

Vancouver  I. 
A.  nanimensis  Ver. 

British  Columbia. 
A.  katherince  Gray. 

Mouth  of  Columbia  River;  Gulf  of  Georgia. 
Leptasterias  leptalea  Ver. 

Southern   Alaska. 
L.  macouni  Ver. 

British  Columbia. 
L.  inequalis  Ver. 

Orca,  Alaska. 
L.  coei  Ver. 

Southern   Alaska. 
L.  vancouveri  (Per.). 

Vancouver  I. 
L.  hcxactis  (St.). 

British  Columbia  to  Monterey,  Calif.;  San  Diego   (?). 
L.  eequalis  (St.). 

British  Columbia;  Puget  Sd.  to  Santa  Barbara. 


SHALLOW- WATER   STARFISHES  341 

Var.  nana  Ver. 

Gulf  of  Georgia  to  California. 
Var.  compacta  Ver. 

British  Columbia  to  Monterey,  Calif. 
L.  epichlora  (Br.)  St. 

Dutch  Harbor;  Aleutian  Is.  to  Vancouver  I. 
Subspecies  alaskensis  Ver. 

Aleutian  Is.  to  Puget  Sd. 
Var.  carinella  Ver. 

Southern  Alaska  to  Dutch  Harbor. 
Var.  siderea  Ver. 

Aleutian  Is.  and  Yakutat  to  Puget  Sd. 
Subspecies  miliaris  Ver. 

Southern  Alaska  to  Queen  Charlotte  Islands. 
Subspecies  plena  Ver. 

Vancouver  I. 
Var.  regularis  Ver. 

Yakutat  to  Sitka,  Alaska. 
Var.  subregularis  Ver. 

Southern   Alaska. 
Var.  subnodulosa  Ver. 

Wrangel,  Alaska. 
Subspecies  pugetana  Ver. 

Puget  Sound. 
L.  ( n  dispar  Ver. 

Dutch  Harbor  to  southern  Alaska. 
Stenasterias  macropora  Ver. 

Southern   Alaska. 
Evasterias  troschelii  (St.). 

Aleutian  Is.  to  Oregon;  abundant  from  Sitka  to  Puget  Sd.  and  Str.  of 

Georgia. 
Var.  rudis  Ver. 

Aleutian  Is.  to  Vancouver  I. 
Var.  densa  Ver. 

British  Columbia;  Vancouver  I. 
Var.  subnodosa  Ver. 

Aleutian  Is.  to  Sitka. 
Var.  alveolata  Ver. 

British  Columbia;  Vancouver  I. 
Var.  parvispina  Ver. 

British  Columbia;  Sitka,  Alaska. 
Evasterias  acanthostoma  Ver. 

Vancouver  I. ;  British  Columbia. 
Orthasterias  columbiana  Ver. 

Yakutat,  Alaska,  to  Victoria,  Vancouver  I. ;  La  Jolla,  Calif. 
O.  biordinata  Ver. 

Departure  Bay,  B.  C 
O.  kcehleri  de  Lor. 

Vancouver  I. 
O.  leptolena  Ver. 

Departure  Bay,  B.  C. 


342  VERRILL 

O.  dawsoni  Ver. 

Queen  Charlotte  Is.,  B.  C. 
O.  (Stylasterias)  forreri  (Lor.)  Ver. 

Monterey  Bay;  Alaska  (?). 
Orthasterias  forreri  forcipulata  Ver. 

Departure  Bay,  B.  C. 
O.  merriami  Ver. 

Glacier  Bay  and  off  Juneau,  Alaska;  Gulf  of  Georgia  (?). 
Distolasterias  chelifera  Ver. 

Vancouver  I.;  Alaska  (?). 
Parasterias  albertensis  Ver. 

British  Columbia. 
Pycnopodia  helianthoides  (Br.)   St 

Aleutian  Is.  and  Yakutat,  Alaska,  to  Monterey  Bay. 
Echinaster  robustus  Ver. 

Departure  Bay,  B.  C. 
Henricia  leviuscula  (St.). 

Bering  I.  and  Aleutian  Is.  to  San    Diego,  Calif. 
Var.  leviuscula  (St.). 

Sitka  to  San  Diego,  Calif. 
Var.  lunula  Ver. 

Sitka  to  Monterey  Bay. 
Var.  attenuata  Dark. 

British  Columbia  and  Vancouver  I.  to  San  Francisco. 
Var.  inequalis  Ver. 

Queen  Charlotte  Is.,  Victoria,  Sitka. 
Subspecies  spiculifera  (Clark). 

Puget  Sd.  to  Bering  Sea. 
Var.  multispina  F. 

Bering  I.  to  Vancouver  I. 
Var.  dyscrita  F. 

Sitka  to  southern  California. 
Var.  annectens  F. 

Puget  Sd.  to  Santa  Barbara,  Calif. 
Var.  spatulifera  Ver. 

Sitka  to  Monterey  Bay;  Santa  Cruz  Is.  (?) 
*H.  sanguinolenta  (Mull.). 

Arctic  Ocean  to  Washington;  0-344  feth. 
*H,  tumida  borealis  Ver. 

Bering  Str.  to  Yakutat  and  Sitka,  Alaska. 
H.  longispina  F. 

S.  Alaska  to  Vancouver  I. ;  41-134  fath. 
H.  aspera  F. 

Bering  Sea  to  Santa  Barbara,  Calif. 
*Solaster  endeca  (L.). 

Arctic  Ocean,  Bering  Sea,  south  to  Sitka;  British  Columbia,  238  fath. 
S.  galaxides  Ver. 

Vancouver  I.  and  Puget  Sd. 
S.  dawsoni  Ver. 

Aleutian  Is.  and  Kuril  Is.  to  Monterey  Bay;  1-229 


SHALLOW-WATER   STARFISHES  343 

S.  constellatus  Ver. 

Puget  Sound. 
*S.  paxillatus  Sladen. 

Bering  Sea,  south  to  Kadiak;  Japan. 
*Crossaster  papposus  (Linne)  Ver. 

Arctic  Ocean,  south  to  Vancouver  I.  and  Puget  Sd. 
Patiria  miniata  (Br.)  Ver. 

Middle  Alaska,  south  to  San  Diego;  Gulf  of  California. 
Pteraster  tesselatus  Ives. 

Bering  I.  to  Puget  Sd. 
P.  tesselatus  hebes  Ver. 

Departure  Bay,  B.  C. 
P.  multispinus  Dark. 

Puget  Sd. 
P.  gracilis  (Clark). 

Puget  Sd. 
*P.  militaris  (Mull.). 

Arctic  Ocean  and  Bering  Sea  to  Aleutian  Is.;  Str.  of  Fuca,  100  fath. 
*Ceramaster  granularis  (Retz.)  V. 

Bering  Sea  to  Puget  Sd. 
C.  patagonicus  (Sladen)   F. 

Bering  Sea;  S.  E.  Alaska;  41-134  fath. 
Mediaster  aqualis  St. 

Alaska  Peninsula  to  Lower  California,  off  Los  Coronados  Is. 
Hippasteria  spinosa  Ver. 

St.  Georges  I.,  Bering  Sea,  and  Kadiak,  south  to  southern  California; 

27-221  fath. 
Dermasterias  imbricata  (Gr.). 

Southern  Alaska  to  Monterey  Bay. 
Var.  valvulifera  Ver. 

Sitka  to  British  Columbia. 
Luidiaster  dawsoni  (Ver.). 

Bering  Sea  to  British  Columbia;  56-159  fath. 
Cteno discus  crispatus  (R.). 

Arctic  Alaska  to  California  in  deep  water. 
Luidia  foliolata  Gr. 

Southern  Alaska  to  San  Diego. 

III.  THE  CALIFORNIAN  FAUNA. 

The  following  list  is  not  thought  to  be  complete.  Probably  many 
more  of  those  in  the  preceding  list  will  be  found  to  extend  into  this 
fauna.  The  two  faunae  blend  together  and  overlap,  so  that  no  very 
definite  division  can  be  made.  This  fauna  is  a  blending  of  the 
preceding  and  following,  with  few  peculiar  species.  The  special 
distribution  of  many  of  the  species  is  given  in  the  preceding  list 
and  not  repeated  here. 


344  VERRILL 

Pisaster  ochraceus  (Br.)  Ag. 
Var.  nodiferus  Ver. 
P.  fissispinus  (St.)  Ver. 

Oregon. 

P.  Wkenii  (St.)  Ver. 
P.  paucispinus  (St.)  Ver. 
fP.  capitatus  (St.)  Ver. 

Monterey  Bay  to  San  Diego. 
P.  giganteus  (St.)  Ver. 

Tomales  Bay,  north  of  San  Francisco. 
P.  brevispinus  (St.)  Ver. 

Crescent  City  to  Monterey,  Calif. 
P.  (?)  gray*  Ver. 

Oregon. 
Asterias  katherince  Gray. 

Mouth  of  Columbia ;  Gulf  of  Georgia. 
Leptasterias  hexactis  (St.)  Ver. 

Monterey  to  Vancouver  I. 
L.  cequalis  (St.)  Ver. 

Vancouver  I.  to  Santa  Barbara,  Calif. 
Var.  compacta  Ver. 

Vancouver  I.  to  Monterey,  Calif. 
Var.  wono  Ver. 

Gulf  of  Georgia  to  Monterey. 
Var.  concinna  Ver. 

Monterey,  Calif. 
Evasterias  troschelii  (St.)  Ver. 

Oregon  and  northward. 
Orthasterias  calif ornica  Ver. 

Near  San  Francisco. 
O.  forreri  (Lor.)  Ver. 

Monterey  and  Santa  Cruz,  Calif.;  Alaska  (?). 
*O.  columbiana  Ver. 

La  Jolla,  Calif.,  to  Yakutat,  Alaska. 
fO.  gonolena  Ver. 

Gulf  of  California ;  Lower  California ;  Santa  Barbara ;  San  Diego,  etc. 
Pycnopodia  helianthoides  (Br.)  St 

Monterey  Bay  northward  to  Kadiak,  Alaska. 
Poraniopsis  infiata  F. 

Oregon  to  San  Diego;  26-159  fath. 
Henricia  leviuscula  (St). 
Var.  lunula  Ver. 
Var.  attenuata  (Qark). 
Var.  dyscrita  F. 
Var.  annectens  F. 
Var.  spatulifera  Ver. 
Patiria  miniata  (Br.)  Ver. 

Yakutat,  Alaska,  south  to  Gulf  of  California. 
Mediaster  cequalis  St 

Alaska,  south  to  Lower  California. 


SHALLOW- WATER   STARFISHES  345 

Hippasteria  spinosa  Ver. 

Kadiak  to  Lower  California ;  27-221  fath. 
Odontaster  crassus  F. 

Monterey  to  San  Diego;  49-284  fath. 
Dermasterias  imbricata  (Gr.). 
Alaska,  south  to  Monterey. 
Astropecten  calif ornicus  F. 

Monterey  Bay  to  Lower  California. 
•fA.  siderealis  Ver. 

Southern  California. 
Bunodaster  ritteri  Ver. 

Californian  coast. 
*Ctenodiscus  crispatus  (Retz.). 

Deep  water ;  California,  north  to  Arctic  Ocean. 
•\Luidia  asthenosoma  F. 

Monterey  to  Lower  California. 
Luidia  foliolata  Gr. 

Alaska  to  San  Diego. 
^L.  ludwigi  F. 

Monterey  to  Santa  Barbara. 

IV.  THE  SOUTH  CALIFORNIAN  FAUNA. 

So  far  as  known,  this  fauna  has  few  species  peculiar  to  it.  The 
species  are  largely  members  of  the  more  northern  faunae  that  extend 
far  south  (indicated  by  an  asterisk),  and  partly  species  of  the  more 
southern  fauna  of  Lower  California  and  the  Gulf  of  California  that 
range  northward  beyond  the  normal  limits  of  that  fauna  (indicated 
by  a  dagger) .  Some  of  the  species,  however,  appear  to  be  peculiar 
to  this  district,  so  far  as  known,  and  others  have  here  their  chief 
development  or  "  center  of  distribution." 

*Pisaster  ochraceus  (Br.). 
Pisaster  capitatus   (St.). 

?  Pisaster  brevispinus  (St.). 
*P.  paucispinus  (St.). 

Santa  Cruz,  Calif. 
*Pisaster  lutkenii  (St.). 
Var.  australis  Ver. 
Var.  exquiseta  (Lor.). 
Santa  Cruz,  Calif. 

^Marthasterias  sertulifera  (Xantus)  V. 
San  Diego  (  ?)  to  Cape  St.  Lucas. 

*Leptasterias  cequalis  (St.).  ... 

Var.  nana  Ver. 
jOrthasterias  gonolena  Ver. 

Santa  Barbara  to  Gulf  of  California. 
O.  forreri  (Lor.)  Ver. 

Santa  Cruz  to  Monterey. 


346  VERRILL 

^Echinaster  tenuisf>inus  Ver. 
San  Diego  to  Panama. 
Poraniopsis  inftata  F. 

San  Diego  to  Oregon. 
*Henricia  leviuscula  (St.). 

San  Diego  north  to  Alaska. 
*Var.  annectens  F. 

Santa  Barbara  to  Puget  Sd. 
*Var.  spatulifera  Ver. 

Santa  Cruz  I.,  northward. 
*H.  aspera  F. 

Santa  Barbara  to  Bering  Sea. 
*Patiria  miniata  (Br.)  Ver. 

San  Diego  and  Gulf  of  California,  north  to  Yakutat,  Alaska. 
*Mediaster  aqualis  St. 

Los  Coronados  Is.,  Lower  California,  and  off  San  Diego,  etc.,  north  to 

Alaska   Peninsula. 
Odontaster  crassus  F. 

San  Diego  to  Monterey  Bay;  43-284  fath. 
*Hippasteria  spinosa  Ver. 

Off  Santa  Catalina  Is.,  Calif.,  in  80  fath.,  north  to  Bering  Sea,  in  121 

fath. 
•\Linckia  Columbia  (Gray). 

Santa  Catalina  Is.  and  San  Diego,  to  Colombia,  S.  A.,  and  Galapagos  Is. 
Astropecten  siderealis  Ver. 

San  Diego  to  San  Pedro. 
A.  calif  or  nicus  F. 

Lower  California  to  Monterey  Bay ;  San  Pedro,  etc. ;  10-244  fath. 
A.  ornatissimus  F. 

Catalina   I.,   San   Pedro,   etc.,   Cerros   Is.   to   Lower   CalifoVnia;  47-207 

fath. 
Luidia  foliolata  (Gr.). 

San  Diego  to  southern  Alaska,  Kassam  Bay;  10-189  fath. 
L.  ludwigi  F. 

Santa  Barbara  and  San  Pedro,  Calif.,  to  Monterey ;  15-50  fath. 
L.  asthenosoma  F. 

Lower  California  to  Monterey;  11-339  fath. 

The  following  species  have  been  erroneously  attributed  to  this 
fauna : 

Asterias  lurida  (Phil.)  =  Cosmasterias  lurida.    See  below,  p.  358. 

Recorded  by  Ives.  A  Chilean  species. 
Asterias  rubens  L. 

Recorded  by  Ives.     European;  not  known  from  the  Pacific  Ocean. 
P hat aria  biserialis  Gray. 

Panamic  fauna. 
Linckia  guildingii  Gray. 

Recorded  by  Ives.    West  Indian. 
Astropecten  cerstedii  Ltk. 

San  Diego  (Ives),  1800.     Panamic  fauna. 


SHALLOW-WATER   STARFISHES  347 

Astropecten  erinaceus  Gray. 

San  Diego  (Ives),  1890.     Panamic  fauna. 
Luidia  brevispina  Liitk. 

San  Diego  (Ives),  1890.     Panamic  fauna. 

V.  SOME    COMPARISONS    WITH    OTHER    FAUN^. 

The  following  remarks  and  lists  are  intended  to  give  only  a  gen- 
eral view  of  the  relations  of  the  Northwest  Coast  faunae  with  those 
of  other  regions. 

As  remarked  in  the  Introduction  and  shown  in  the  preceding  lists, 
a  considerable  percentage  of  the  starfishes  of  the  Beringian  fauna  are 
circumpolar. 

The  total  number  of  species  and  subspecies  listed  in  the  Beringian 
fauna  is  forty-four.  Of  these  fifteen,  or  thirty-three  per  cent,  are 
known  to  be  circumpolar.  Eight  species,  or  about  eighteen  per  cent, 
appear  to  belong  more  properly  to  the  Columbia-Alaskan  fauna, 
while  many  of  the  remaining  species  extend  southward  into  the  latter. 
Six  of  the  remainder  reach  Siberia,  and  four  species  reach  Japan. 

About  sixteen  species  are  either  peculiar  to  the  fauna  or  have  there 
their  principal  development,  so  far  as  known.  Of  these,  several  are 
so  closely  allied  to  North  Atlantic  and  Arctic  species  that  they  must 
be  considered  as  of  arctic  or  subarctic  origin.  Such  are  Asterias 
multiclava;  A.  polythela;  Leptasterias  arctica;  Henricia  sanguino- 
lenta  var.  rudis;  H.  tumida  borealis;  H.  tumida;  H.  arctica;  Solaster 
dawsoni  arctica;  Pteraster  marsippus;  Pteraster  obscures  octaster; 
Pterasterides  aporus;  Leptychaster  pacificus. 

It  is  not  improbable  that  most  of  these,  as  well  as  various  species 
now  circumpolar  in  range,  may  have  originated  in  Bering  Sea  or 
adjacent  waters. 

A  few  species  are  identical  with,  or  more  closely  allied  to,  northern 
Japanese  or  Asiatic  species  than  to  those  of  any  other  fauna.  These 
are  Allasterias  rathbuni,  and  subspecies  nortonensis;  A.  anomala; 
Solaster  paxillatus,  Patiria  miniata,  and  Glyphaster  anomalus.  Such 
species  may  have  had  their  origin  on  the  Asiatic  coast,  for  the  oceanic 
currents  are  mainly  from  that  direction. 

When  compared  with  the  Antarctic  species,  we  find  very  little 
affinity,  except  in  the  cases  of  a  few  very  widely  distributed  deep- 
water  genera  and  species,  such  as  Ceramaster  patagonicus,  Ctenodis- 
cus  crispatus.  These  are  of  doubtful  origin,  having  related  species 
in  many  regions.  See  below. 


348  VERRILL 

VI.  RELATIONS  OF  THE  COLUMBIA-ALASKAN  AND 
CALIFORNIAN   FAUNAE. 

The  former  includes  a  total  of  eighty-five  species  and  named 
varieties;  the  latter  includes  forty  altogether.  For  my  present 
purpose  these  may  best  be  considered  collectively,  as  they  have  many 
species  in  common.  The  two  lists  include  ninety-nine  species,  sub- 
species, and  varieties.  Of  these  only  sixteen  species  and  varieties 
are  not  known  to  occur  in  the  Columbia- Alaskan  fauna,  while  twenty- 
four  are  common  to  both. 

Of  the  ninety-nine  forms,  only  about  eight  are  of  Arctic  origin 
(indicated  by  an  asterisk  in  the  list),  leaving  nine-one  that  may  be 
considered  as  characteristic  of  the  region.  Of  this  number  a  few  are 
more  or  less  related  to  Arctic  species  generically,  or  have  there  their 
nearest  allies. 

Among  these  are  Asterias  katherina;  A.  victoriana;  Leptasterias 
leptalea;  Solaster  galaxides;  Hippasteria  spinosa. 

A  few  have  their  nearest  known  allies  on  the  Atlantic  coast  of  the 
United  States.  Among  these  are  Orthasterias  columbiana,  allied  to 
O.  tanneri  of  the  Atlantic ;  Mediaster  aqualis,  allied  to  M .  bairdii. 

A  considerable  number  belong  to  genera  widely  distributed  in  all 
the  warmer  or  temperate  seas.  Among  these  are  the  various  forms  of 
Henricia  leviuscula;  H.  longispina;  H.  aspera;  Solaster  stimpsoni; 
S.  dawsoni;  S.  constellates;  Pteraster  tesselatus;  P.  multispinus;  P. 
gracilis;  one  species  of  Astropecten,  and  three  of  Luidia. 

The  species  of  Astropecten  and  Luidia  are  probably  of  tropical 
origin  and  their  nearest  allies  apparently  still  exist  in  the  Panamic 
fauna. 

Many  species  and  several  genera  of  this  region  do  not  appear  to 
have  any  very  closely  allied  species  in  other  seas.  Among  these  are 
the  following:  The  genus  Pisaster,  with  its  eleven  species  and  sub- 
species; Leptasterias  epichlora,  with  its  ten  subspecies  and  named 
varieties,  none  of  which  seem  closely  related  to  Atlantic  species ;  L. 
coei;  L.  hexactis;  L.  aqualis,  with  three  varieties;  Evasterias  tro- 
schelii,  with  five  varieties ;  E.  acanthostoma;  eight  species  of  Orth- 
asterias; Parasterias  albertensis;  Stenasterias  macropora;  Pycno- 
podia  helianthoides ;  Dermasterias  imbricata;  Bunodaster  ritteri. 

The  last  five  are  the  only  known  representatives  of  their  genera. 

A  few  species  seem  to  be  allied  to  some  of  those  of  the  coasts  of 
Peru  and  Chile,  without  any  allies  in  the  intervening  tropical  waters. 
Thus,  Patiria  miniata  is  not  distantly  related  to  P.  chilensis,  with  no 
species  of  the  same  genus  known  between  Lower  California  and 


SHALLOW-WATER   STARFISHES  349 

Peru,  nor  on  the  Atlantic  coasts  of  America.  Yet  a  much  more 
closely  allied  species,  P.  pectinifera  (Miiller  and  Troschel),  is  found 
on  the  coasts  of  Japan,  extending  north  at  least  as  far  as  Vladivostok. 
Thus,  the  North  American  species  is  most  likely  a  derivative  of  the 
Japanese  stock.  The  restricted  genus,  itself,  is  widely  distributed  in 
the  Indian  and  Pacific  oceans,  from  Cape  Good  Hope  to  Australia 
and  Japan,  but  the  Japanese  species  referred  to  is  decidedly  nearer  to 
P.  miniata  than  any  of  the  others.1  All  the  species  of  the  genus  are 
littoral  and  shallow-water  forms.  This  renders  their  wide  distribu- 
tion more  remarkable.  Poraniopsis  inflata  Fisher  is  allied  to  P. 
echinaster  Perrier,  of  Patagonia,  etc.  Odontaster  crassus  seems  to  be 
related  to  the  several  forms  of  the  same  genus  found  off  Patagonia 
and  Cape  Horn,  but  perhaps  no  more  so  than  to  the  three  species 
found  off  the  Atlantic  coast  of  the  United  States,  especially  O. 
robustus  Verrill  and  O.  hispidus  Verrill.  Ceramaster  patagonicus  is 
thought  by  Professor  Fisher  to  have  a  continuous  range,  in  the  deep 
sea,  from  Patagonia  to  Alaska  and  Japan.  The  same,  he  thinks,  may 
be  true  of  Ctenodiscus  crispatus.  I  have  not  personally  studied  Ant- 
arctic specimens  of  these  species.  Luidiaster  dawsoni  belongs  to  a 
deep-sea  genus,  widely  distributed. 

The  preceding  brief  analysis  of  the  fauna!  lists  indicates  pretty 
clearly  that  much  the  larger  part  of  the  starfishes  inhabiting  shallow 
waters  of  the  North  Pacific  coast,  from  middle  California  to  Southern 
Alaska,  originated  on  that  coast  and  have  received,  at  most,  only  a 
few  additions  from  the  Arctic  Ocean  and  from  the  Panamic  and 
South  American  faunae,  in  modern  (geologic)  times. 

About  one-half  (forty-nine)  of  the  total  number  of  species  and 
varieties  in  the  lists  belong  to  the  family  Asteriidae. 

VII.  RELATIONS  OF  THE  SOUTH  CALIFORNIAN  FAUNA. 
The  list  above  includes  twenty-nine  species  and  varieties.  Of  these, 
twenty-two  species  occur  farther  north.  Of  the  remaining  six  species, 
three  are  known  to  occur  in  the  Panamic  fauna,  and  probably  find 
here  their  northern  limits,  viz.,  Marthasterias  sertulifera;  Echinaster 
tenuispinus;  Linckia  Columbia.  These  belong  to  genera  widely  dis- 
tributed in  tropical  seas.  The  remaining  three,  at  present,  seem  to 
belong  particularly  to  this  fauna,  viz.,  Astropecten  siderealis;  A. 
ornatissimus  ;  Orthasterias  gonolena. 

1  Professor  Fisher,  op.  cit.  iQiifr,  p.  258,  has  given  a  rery  useful  table  of  their 
differences. 


350  VERRILL 

The  two  species  of  Astropecten  are  closely  allied  to  Panamic  spe- 
cies. The  Orthasterias  is  allied  to  O.  calif  arnica  and  others  from 
farther  north,  but  it  ranges  to  the  Gulf  of  California. 

This  faunal  district,  as  now  known,  does  not  show  any  special 
peculiarities  of  its  own.  It  is  a  meeting-ground,  so  to  speak,  between 
the  Panamic  and  the  Californian  faunae.  Yet  the  list  of  species 
given  above  is  doubtless  very  incomplete. 

It  is  notable  that,  while  twelve  species  and  varieties  of  Asteriinae 
reach  this  fauna,  only  two  extend  southward  to  Cape  St.  Lucas. 
From  thence  southward,  on  the  whole  extent  of  the  coast  to  Ecuador, 
no  member  of  this  subfamily  is  known  to  occur  in  shallow  water. 

From  off  Mexico,  the  Galapagos  Islands,  and  Panama,  in  moder- 
ately deep  water,  Ludwig  has  described  seven  species,  which  he  refers 
to  Stolasterias,  Sporasterias,  and  Hydrasterias.  One  of  these  appears 
to  belong  to  Coscinasterias,  viz.,  C.  alexandri  (Ludwig,  as  Stolas- 
terias, p.  221,  pi.  xxxn,  figs.  190-193).  It  is  from  the  Gulf  of 
Panama,  in  95  meters  to  384  meters. 

Another,  which  appears  to  belong  to  Stylasterias,  viz.,  S.  robusta 
(Ludwig,  as  Stolasterias,  p.  228),  is  from  off  the  Galapagos,  in 
704  meters. 

The  three  species  referred  to  Sporasterias  do  not  seem  to  belong  to 
that  genus  as  herein  restricted. 

S.  mariana  is  from  off  Las  Tres  Marias  Islands,  in  1244  meters ; 
.S".  cocosana  is  from  245  meters,  off  Cocos  Island ;  S\  galapagensis  is 
from  off  Chatham  Island,  in  704  fathoms. 

The  dorsal  plates  and  connective  ossicles  are  broad,  imbricated,  and 
have  a  regularly  stichasterial  arrangement  in  the  last  two  species, 
which  are  otherwise  very  distinct.  , 

6".  mariana  has  five  rows  of  dorsal  plates  and  no  visible  interactinals. 
It  may  belong  to  Distolasterias. 

Hydrasterias  diomedecs  is  from  155  meters  and  121  meters,  off 
Panama  and  Cocos  Island. 

None  of  these  species  are  nearly  allied  to  any  of  the  shallow  water 
species  of  California. 

The  subfamily  Asteriinae  is  again  abundantly  represented  by  shal- 
low-water species  on  both  coasts  of  southern  South  America  from 
Chile  and  northern  Patagonia  to  Cape  Horn  and  the  Antarctic 
Islands.  Yet  nearly  all  the  species  of  these  regions  belong  to  general 
not  found  in  the  North  Pacific  and  North  Atlantic. 


SHALLOW- WATER   STARFISHES  351 

VIII.  PATAGONIAN,  FUEGIAN,  AND  ANTARCTIC  SHALLOW- 
WATER  STARFISHES  CONTRASTED  WITH  THOSE  OF 
THE  NORTH  PACIFIC  COAST.1 

As  already  remarked,  the  Antarctic  starfishes  are  abundant  and 
diversified,  especially  those  of  the  subfamily  Asteriinae.  Most  of  the 
latter,  found  in  shallow  water,  belong  to  genera  peculiar  to  the 
southern  hemisphere. 

One  notable  peculiarity  is  found  in  the  large  proportion  of  species 
that  carry  and  incubate  their  eggs  and  young,  being  brooders,  or 
padophoric  species.  This  peculiarity  extends,  also,  to  many  of  the 
Antarctic  Echinoidea,  Ophiuroidea,  and  Holothurioidea. 

The  genera  of  starfishes  of  the  family  Asteriidae,  having  this  habit, 
differ,  also,  in  anatomical  details  and  in  the  position  of  the  genital 
pores  from  those  that  produce  minute  eggs  that  develop  into  free- 
swimming  larvae,  as  is  commonly  the  case  with  northern  genera. 

Most  of  the  pcedophoric  Asteriidae  carry  .their  young  in  clusters 
under  and  around  the  mouth,  as  in  Leptasterias  of  the  north,  but 
Stichaster  nutrix  Studer  is  described  as  carrying  them  in  pouch-like 
diverticula  of  the  stomach,  at  least  in  part. 

This  padophoric  habit  is  not  confined  to  shallow-water  and  littoral 
species,  for  it  has  been  observed  in  some  species  from  considerable 
depths,  such  as  Anastfrias  belgicce  Ludwig,  and  A.  chirophora  Lud- 
wig,  taken  in  450  to  560  meters,  in  the  Antarctic  Ocean  (see  below). 
In  these  species  the  young  are  attached  together  in  large  groups,  like 
clusters  of  grapes,  by  means  of  their  adoral  peduncles  and  a  central 
stalk. 

Although  many  of  the  Antarctic  species  agree  with  Leptasterias 
of  the  north,  in  this  habit  of  carrying  their  young,  and  some  of  them, 
also,  in  general  appearance,  they  do  not  appear  to  be  nearly  allied.1 
Most  of  them  are  regularly  monacanthid,  while  Leptasterias  is  never 
truly  monacanthid.  Some  are  diplacanthid,  as  in  the  case  of  Podas- 
terias  steineni  and  P.  liitkenii  (see  below,  p.  361). 

In  the  case  of  Leptychaster  kerguelenensis  Smith,  the  eggs  and 
young  are  carried  on  the  back,  under  the  protection  of  the  projecting 
spinules  of  the  paxillae,  which  is  more  analogous  to  the  method  in 
Pterasteridae,  in  which  the  supraspinal  membrane  gives  additional 
protection,  and  for  a  longer  period  of  development. 

1  In  the  following  review  I  have  omitted  some  imperfectly  described  forms 
and  all  the  abyssal  species. 

*It  may  be  safely  assumed,  I  believe,  that  all  species  that  carry  their  eggs 
and  young  around  the  mouth,  also  have  their  genital  pores  on  the  ventral  side, 
and  therefore  cannot  be  congeneric  with  those  that  have  them  dorsal,  as  in 
typical  Asterias. 


352  VERRILL 

Another  striking  peculiarity  of  the  Asteriinae  of  this  region,  and  of 
the  Antarctic  Ocean  generally,  is  the  large  percentage  of  species  that 
have  the  skeletal  plates,  and  sometimes  the  spines,  covered  with  a 
thick  and  soft,  but  rather  tough,  dermis,  which  is  usually  canalicu- 
lated,  and  often  entirely  conceals  the  plates  in  alcoholic  preparations, 
or  even  in  dry  ones.1  Even  the  spines,  if  small,  may  be  submerged  in 
the  dermis,  or  appear  as  mere  papillae. 

In  some  cases  such  starfishes  have  the  skeletal  plates,  more  or  less 
aborted,  especially  the  dorsal  ones,  but  sometimes  even  the  upper 
marginals,  as  in  Anasterias,  Padasterias,  and  Adelasterias. 

FORCIPULOSA. 

No  doubt  the  most  peculiar  and  remarkable  species,  pertaining  to 
these  southern  waters,  is  the  Labidiaster  radiosus  Liitken,  belonging 
to  the  Brisingidae. 

It  has  a  small  disk  with  from  twenty-six  to  forty-two  rays,  the 
number  increasing  with  age,  and  grows  to  a  large  size.  Unlike  the 
other  forms  of  Brisingidae,  it  lives  in  shallow  water,  as  well  as  at  a 
considerable  depths.  It  is  found  on  both  coasts  of  Patagonia  and  off 
Cape  Horn,  etc. 

The  only  other  described  species  is  L.  annulatus  Sladen.  It  oc- 
curred off  Kerguelen  Island  and  Heard  Island,  in  75  to  150  fathoms, 
and  in  the  Arafura  Sea  (  ?)  in  800  fathoms.  It  has  forty  to  forty-five 
long,  slender  rays.  It  differs  considerably  in  structure  from  L.  radio- 
sus, especially  in  having  the  dorsal  and  superomarginal  plates  nearly 
abortive  distally,  on  the  rays,  beyond  the  genital  regions.  This  last 
character,  with  others,  should  require  a  generic  separation. 

Therefore  I  propose  for  it  the  new  generic  name,  Labidastrella, 
with  L.  annulata  as  its  type. 

Aside  from  Labidiaster,  the  most  characteristic  species  belong  to 
the  Asteriidae  and  Ganeriidae.  There  are  several  genera  of  Asteriidae 
that  do  not  occur  in  northern  seas,  while  most  of  the  northern  genera 
are  entirely  lacking.  Thus  there  are  no  species  of  true  Asterias, 
Pisaster,  Orthasterias,  Evasterias,  Urasterias,  Allasterias.  It  is 
doubtful  if  there  is  any  true  Leptasterias,  though  some  antarctic 

1  The  apparent  thickness  of  the  dermis  depends  much  on  the  mode  of  pres- 
ervation, which  should  be  stated  in  descriptions.  It  is  hardened  and  con- 
tracted by  strong  alcohol,  and  still  more  when  dried,  especially  after  being  in 
alcohol.  In  alcohol  of  about  70  per  cent  it  is  more  naturally  preserved.  In 
very  weak  alcohol  and  in  formalin  it  may  swell  up  and  become  soft.  Some 
of  the  difficulties  in  the  identification  of  such  starfishes  are  thus  due  to  the 
mode  of  preservation.  In  more  cases  it  is  due  to  the  negligence  of  the  describer 
in  not  cleaning  and  describing  the  skeletal  plates  and  pedicellariae. 


SHALLOW-WATER   STARFISHES  353 

species  have  been  described  that  closely  resemble  that  genus,  and  may 
possibly  belong  to  it. 

On  the  other  hand,  there  are  found  in  the  Patagonian  fauna,  and 
farther  south,  numerous  species  and  varieties  of  monacanthid  star- 
fishes, most  of  which  are  known  to  carry  their  eggs  and  young  (pcedo- 
phoric)  and  to  have,  in  those  species  dissected,  ventral  genital  pores. 
These  have  been  referred  to  the  genera  Anasterias  Perrier,  1875 ; 
Calvasterias  Perrier,  1875;  Sporasterias  Perrier,  1894;  Pcedasterias 
Verrill,  gen.  nov. 

Another  group  of  pcedophoric  species  includes  several  diplacanthid 
species,  belonging  to  the  genera  Podasterias  Perrier  and  Granaster 
Perrier.  None  of  these  are  known  in  the  northern  hemisphere. 

The  genera  Meyenaster  Verrill,  May,  1913  (monacanthid),  and 
Cosmasterias  Sladen,  1889  (diplacanthid),  also  characteristic  of  this 
region,  have  not  yet  been  observed  to  carry  their  young. 

The  most  common  and  abundant  littoral  and  shallow-water  star- 
fishes of  both  coasts  of  Terra  del  Fuego  and  Magellan  Strait  belong 
to  a  monacanthid  generic  group  with  a  reticulated  dorsal  skeleton, 
for  which  Sporasterias  Perrier,  1894,  seems  to  be  the  earliest  tenable 
name,  with  S.  rugispina  (Stimpson,  1862,  as  Asterias)  for  its  type. 
This  group,  which  we  may  also  designate  as  the  rugispina — or  ant- 
arctica — group,  has  commonly  been  called  Anasterias  Perrier.  The 
latter  was  originally  based  solely  on  A.  minuta  Perrier,  a  small,  doubt- 
less very  young  starfish,  with  a  thick  dermis,  rudimentary  dorsal 
skeleton,  and  few  dorsal  spines,  and  thus  quite  unlike  the  group 
that  we  call  Sporasterias.  (See  Anasterias,  below,  p.  354.) 

F.  Leipoldt  (op.  cit.,  1895,  p.  563)  proposed  to  unite  nearly  all  the 
then  known  monacanthid  starfishes  of  the  Antarctic  seas  into  one 
comprehensive  and  exceedingly  variable  species.  This  would  be  a 
very  convenient  arrangement,  perhaps,  if  it  would  only  prove  true. 
No  doubt,  however,  many  of  the  nominal  species  of  this  group  are 
synonymous ;  and  the  species  are  certainly  variable,  as  are  all  others 
that  carry  and  incubate  their  eggs  and  young. 

Among  the  forms  united  by  Leipoldt,  under  the  name  Asterias 
rugispina,  the  following  belong  apparently  to  Sporasterias,  having  a 
reticulated  dorsal  skeleton  and  many  spines,  not  submerged  in  a  thick 
dermis :  Sporasterias  antarcticd '(Liitken,  1856)  ; S.  rugispina  (Stimp- 
son, 1862,  as  Asterias)  ;  5".  varia  (Phil.,  1870)  ;  S.  rupicola  (Verrill, 
1876,  as  Asterias) ;  S.  perrieri  (Smith,  1876,  six-rayed,  as  Asterias, 
non  A.  perrieri  Studer,  1884)  ;  5.  cunninghami  (Perrier,  1875,  as 
Asterias)  ;  5\  spirabilis  (Bell,  1881,  as  Asterias)  ;  S.  hyadesi  (Per- 

24 


354  VERRILL 

rier,  1886,  as  Asterias).  Of  these,  S.  rugispina  appears  to  be  dis- 
tinct by  reason  of  its  large,  wide,  obtuse  minor  pedicellariae,  and 
few,  capitate  and  sulcate  dorsal  spines.  6*.  perrieri  Smith,  from 
Kerguelen  Island,  is  regularly  six-rayed;  S.  rupicola  Verrill,  from 
the  same  place,  a  smaller,  five-rayed  form,  with  many  spines  and 
few  and  minute  pedicellariae,  is  apparently  distinct.  (See  below,  p. 
356.)  All  the  others  may  well  belong  to  a  single  species,  for  which 
S.  antarctica  (Liitken)  seems  to  be  the  correct  name. 

AN  ASTERIAS  (Perrier)  restricted. 

In  addition  to  all  the  above  forms,  Leipoldt  includes  in  the  same 
composite  species  the  following,  which  have  a  rudimentary  or  partly 
abortive  dorsal  skeleton;  few  small  imbedded  spinules;  and  a  thick 
dermis  that  conceals  the  plates,  and  largely  the  spinules.  These 
constitute,  with  others,  the  restricted  genus  Anasterias  (Lysasterias 
Fisher)  :  Anasterias  minuta  Perrier,  1875  (type)  ;  A.  perrieri  (Stu- 
der),  1884  (non  Smith)  ;  A.  verrillii  (Bell,  1881,  as  Asterias).  Also 
Calvasterias  stolidota  Sladen. 

Fisher  (1908,  p.  88)  proposed  the  name  Lysasterias  to  replace 
Anasterias  Ludwig,  with  Asterias  perrieri  Studer,  of  South  Georgia, 
as  the  type,  assuming  that  A.  minuta  belongs  to  a  distinct  genus  and 
is  identical  with  S.  rugispina,  etc.  However,  the  original  description 
by  Perrier  shows  that  it  has  only  isolated  dorsal  plates,  covered  with 
a  thick  dermis ;  few  imbedded  dorsal  spinules ;  numerous  pedicellariae, 
and  most  of  the  other  characters  as  in  A.  perrieri  Studer  and  A. 
studeri  Perrier.  Therefore,  it  is  most  likely  the  young  of  one  of  these 
or  of  some  other  closely  allied  species  of  the  same  group,  and  not  the 
young  of  rugispina,  nor  of  spirabilis,  as  has  been  thought,  for  the 
young  of  that  group,  when  of  similar  size,  have  a  reticulated  dorsal 
skeleton,  abundance  of  spines,  and  the  dermis  not  thick  enough  to 
conceal  the  plates.  Hence,  I  consider  Lysasterias  Fisher  a  synonym 
of  Anasterias. 

Since  the  specific  name,  Asterias  perrieri  Studer  (1884),  was  pre- 
occupied by  A.  perrieri  Smith,  1876,  it  requires  a  new  name.  There- 
fore, I  propose  to  call  it  ANASTERIAS  LYSASTERIA. 

To  the  genus  Anasterias  (restr.)  Ludwig  (1905,  p.  42)  added, 
besides  the  two  species  above  named,  three  new  species  from  deeper 
water,  southwest  of  Cape  Horn.  The  described  species  are  as 
follows : 


SHALLOW-WATER   STARFISHES  355 

A.  minuta  Per.,  1875. 

Patagonian  and  Fuegian;  shallow-water. 
A.  lysasteria  Ver.,  nom.  nov.  =  A.  perrieri  (Std.  non  Smith). 

South  Georgia,  cast  ashore;  also  Fuegian  (t.  Perrier). 
A.  studeri  (Per.),  1891. 

Falkland  Is.,  320  meters ;  carried  young. 
A.  verrillii  (Bell,  1881,  p.  513,  pi-  XLVII,  fig.  3,  as  Asterias). 

Fuegian ;  Gregory  Bay,  etc.,  littoral. 
A.  lactea  Lud.  (1903,  p.  50). 

About  S.  lat.  71°,  in  450  meters. 
A.  belgica  Lud.  (1903,  p.  51). 

About  S.  lat.  70°,  in  560  meters ;  carried  young. 
A.  tenera  (Kcehler,  1905). 

Wandel  I. ;  carried  young. 

The  attached  young  of  A.  belgicce  were  described  by  Ludwig,  1903, 
p.  53,  pi.  vi,  figs.  61-65  ;  pi.  vii,  figs.  66,  67. 

;        For  the  following  species,  placed  in  Anasterias  by  Ludwig,  I  pro- 
pose to  establish  a  new  genus : 

Genus  Paedasterias  Verrill,  nov. 
Type  A.  chirophora  (Ludwig,  op.  cit.,  1903,  p.  43,  plates  v,  vi,  vn). 

This  is  monacanthid  and  incubates  its  young,  and  therefore  prob- 
ably has  ventral  genital  pores.    Its  skeleton  is  more  reduced  than  in     x^_. 
Anasterias,  the  upper  marginal  plates  being  absent,  except  as  rudi- 
ments distally.    It  is  covered  by  a  thick  dermis  and  bears  large,  wide, 
felipedal  pedicellariae. 

It  is  found  at  from  450  to  560  meters,  in  .the  Antarctic  Ocean,  south- 
west of  Cape  Horn,  about  S.  lat.  70°  to  71°. 

Ludwig  has  carefully  described  and  figured  the  attached  young. 

Genus  Sporasterias  Perrier. 
Type,  Asterias  rugispina  Stimpson. 

Sporasterias  PERRIER,  1896,  p.  55. 
Anasterias  (Pars}  PERRIER,  1891  (non  1875). 
Sporasterias  LUDWIG,  1905. 

This  is  a  monacanthid  genus  with  a  reticulated  dorsal  skeleton,  not 
replaced  by  a  thick  dermis,  bearing  numerous  small,  scattered  spines, 
and  usually  numerous  pedicellariae  of  both  kinds. 

Interactinal  spines  are  usually  absent;  sometimes  there  is  an 
imperfect  row.  Inferomarginal  spines  are  usually  two  to  a  plate; 
superomarginal  ones  usually  one  to  a  plate;  interactinal  papulae 
usually  solitary,  enlarged  proximally ;  jaws  and  adoral  ridge  narrow, 
elongated. 


VERRILL 

SPORASTERIAS  PERRIERI  (Smith). 

Asterias  perrieri  SMITH,  Ann.  and  Mag.  Nat.  Hist.,  ser.  4,  xvn,  p.  106,  1876 
(non  Studer,  1884;  non  Anasterias  perrieri  Ludwig,  1905). 

This  is  a  true  Sporasterias,  with  well  developed  skeleton  and  numer- 
ous spines.  It  is  reguarly  six-rayed.  Smith  states  that  his  largest 
specimen  carried  hundreds  of  attached  young,  all  of  which  were  six- 
rayed.  It  appears  to  be  clearly  distinct  from  the  rugispina-antarctica 
group. 

The  type  was  from  Kerguelen  Island,  in  5  to  10  fathoms. 

SPORASTERIAS  RUPICOLA  Verrill. 

Asterias  rupicola  VERRILL,  op.  cit.,  1876,  p.  74. 

This  is  a  small,  short-rayed  form.  The  types,  which  were  from 
Kerguelen  Island  shore,  have  the  following  characters  when  dry : 

The  dorsal  skeleton  is  pretty  well  developed,  openly  reticulated, 
with  numerous  moderately  large  papular  areas,  evenly  distributed, 
larger  on  the  basal  part  of  the  rays,  mostly  with  two  to  four  or  five 
papulae;  single  ones  occur  distally.  The  dorsal  plates  are  not  con- 
cealed by  the  dermis ;  the  dorsal  spines  are  evenly  distributed,  short, 
clavate,  subequal,  with  no  evident  larger  median  row,  one  to  three 
on  the  small  nodal  plates. 

The  superomarginal  spines  form  a  distinct  simple  row,  one  to  a 
plate.  The  plates  being  stout  and  imbricated.  The  inferomarginal 
plates  mostly  bear  two  larger  and  longer  clavate  spines.  A  wide  lane 
between  the  upper  and  lower  marginal  rows,  broadening  proximally, 
has  a  row  of  large  papular  areas,  with  three  to  five  papulae  separated 
by  stout  descending  apophyses  of  the  superomarginal  plates,  some 
of  which  carry  a  single  small  spinule. 

A  short  interactinal  row  of  small  plates  exists  proximally,  some  of 
them  bearing  a  single  spine.  Between  these  is  a  row  of  single  papulae ; 
the  proximal  ones  are  often  larger  than  the  spines  and  inflated,  as  in 
other  allied  species. 

The  adambulacral  spines  form  a  single  regular  row;  they  are 
smaller  and  more  slender  than  the  inferomarginals,  and  are  distinctly 
clavate.  The  interradial  axils  are  covered  with  a  thicker  canaliculated 
dermis  without  spines.  The  jaws  are  narrow. 

Major  pedicellariae  a*e  few  and  small;  some  along  the  furrow 
margins  and  on  the  oral  spines  are  thick,  ovate  or  blunt-lanceolate ;  a 
few  smaller,  more  acute  ones  are  on  dorsal  spines.  Minor  pedicellariae 
appear  to  be  lacking  in  the  several  types.  They  are  abundant  in 
6".  rugispina  (Stimpson)  and  most  of  the  other  allied  forms,  and  there 
are  other  notable  differences  in  the  spines  and  skeleton. 


SHALLOW-WATER   STARFISHES  357 

Genus  Calvasterias  Perrier. 
Type,  C.  asterinoides  Perrier;  Sladen,  1889,  p.  589. 
Calvasterias  PERSIES,  op.  cit.,  1875,  p.  348. 

This  is  another  genus  in  which  the  skeleton,  and  also,  largely,  the 
spines,  are  covered  with  a  thick,  soft  dermis,  entirely  concealing  the 
plates. 

It  is  monacanthid.  An  imperfect  spiniferous  interactinal  row  of 
plates  is  present ;  usually  there  are  two  rows  of  inf  eromarginal  spines 
and  a  single  row  of  superomarginals.  Position  of  genital  pores 
unknown ;  nor  is  it  known  whether  the  young  are  incubated. 

The  dorsal  skeleton  in  the  type  consists  of  wide,  lobed  or  substellate, 
imbricated  plates,  overlapping  in  both  directions  by  their  lobes,  as  in 
Stichaster.  Descriptions  and  figures  of  other  species  do  not  show  the 
character  of  the  plates,  and  some  may  not  be  congeneric,  if  the  plates 
prove  to  be  abortive  or  rudimentary. 

In  the  type  the  dorsal  spines  are  few,  short,  capitate,  nearly  con- 
cealed by  the  dermis  and  an  abundance  of  papulae  of  similar  size. 

The  type  was  supposed  to  have  been  from  Torres  Strait,  but  Sladen 
records  the  species  from  the  Falkland  Islands. 

CALVASTERIAS   (?)   STOLIDOTA  Sladen. 
C.  stolidota  SLADEN,  1889,  p.  590,  pi.  ci,  figs.  3,  4;  pi.  an,  figs,  n,  12. 

This  has  longer  rays  than  the  type,  with  more  evident,  scattered 
dorsal  spines,  which  are  short,  rough,  capitate.  The  skeleton  was  not 
described,  and  so  it  may  not  be  a  true  Calvasterias.  Leipoldt  united  it 
to  Sporasterias  rugispina,  with  doubt. 

This  is  from  both  coasts  of  Patagonia  and  the  Falkland  Islands, 
in  5  to  10  fathoms. 

MEYENASTER  GELATINOSUS   (Meyen,   1834,  as  Asterias). 

Asterias  rustica  GRAY,  1840,  p.  179;  Synopsis,  p.  i,  1866. 
Asterias  gelatinosa  CLARK,  op.  cit.,  1910,  p.  337,  pi.  vi,  fig.  2. 
Meyenaster  gelatinosus  VERBILL,  Amer.  Journ.   Science,  xxxv,  May,   1913, 
p.  348.    See  also  Introduction,  above,  p.  54,  for  generic  characters. 

This  is  a  large,  and  a  very  characteristic  species  on  the  west  coast 
of  Chile,  from  Iquique  southward.  It  grows  to  the  diameter  of 
20  inches  or  more.  It  is  monacanthid  and  usually  six-rayed,  yet  five- 
rayed  and  seven-rayed  specimens  are  frequently  found. 

The  only  other  species  of  .the  genus,  known  to  me,  is  supposed  to  be 
from  the  Society  Islands  (No.  1427,  Coll.  Mus.  Comp.  Zool.).  That 


VERRILL 

is  a  large,  five-rayed  species,  similar  as  to  spinulation  and  thick 
dermis,  but  with  very  unlike  pedicellariae. 

Its  major  pedicellariae  are  remarkably  slender,  numerous,  part  of 
them  large,  narrow-lanceolate  and  acute.  The  minor  pedicellariae 
are  minute  and  very  abundant,  both  on  the  dermis  and  on  thick  sheaths 
around  the  sulcated  dorsal  spines. 

This  genus  somewhat  resembles  Marthasterias. 

*» 

Genus  Cosmasterias  Sladen,  1889. 
Type,  Asterias  sulcifera  Perrier,  1869,  =  C.  lurida  (Phil.). 
Diplasterias  (pars)  PERKIER,  1891. 

This  is  also  a  genus  apparently  peculiar  to  the  Patagonian  and 
Fuegian  region,  where  one  species  is  abundant  in  shallow  water  and 
often  at  low  tide. 

It  is  diplacanthid ;  jaw-spines  and  adorals  are  elongated ;  the  dorsal 
plates  form  several  longitudinal  bands  or  rows,  and  in  the  adult  each 
plate  bears  a  group,  of  short,  unequal  spines.  There  is  at  least  one 
row  of  interactinal  spines;  often  three  or  more  rows,  when  adult. 
Large  unguiculate  or  felipedal  pedicellarise  are  found,  above  and 
below.  Neither  of  the  species  has  been  observed  to  carry  the  young. 

COSMASTERIAS  LURIDA  (Phil.,  1858)  Ludwig,  1905. 
Cosmasterias  sulcifera   (PERRIER,   1869,  as  Asterias)    Sladen,  1889.     To  this 
species   Leipoldt,   1895,   unites  the   following:    Asteracanthion   clavatum 
Phil. ;  A.  fulvum  Phil. ;  A.  spectabile  Phil. ;  A.  mite  Phil. ;  and  Stichaster 
polygrammus  Sladen,  1887,  245  fathoms. 

The  species  last  named  seems  to  me  clearly  distinct.  (See  below, 
p.  360.) 

To  the  synonyms  of  this  species  it  will  probably  be  necessary  to  add 
Asterias  obtusispinosa  Bell,  1881,  p.  92,  pi.  ix,  fig.  3 ;  and  A.  neglecta 
Bell,  op.  cit.,  p.  94,  pi.  ix,  fig.  4.  Both  are  Patagonian ;  the  former 
from  Sandy  Point,  9  to  10  fathoms.  C.  alba  (Bell),  loc.  cit.,  p.  92, 
pi.  ix,  fig.  2,  also  from  Sandy  Point,  may  not  be  distinct,  though  it  has 
somewhat  longer  spines. 

C.  sulcifera  has  been  considered  the  type  of  Cosmasterias  Sladen 
and  Diplasterias  Perrier.  (See,  also,  Introduction,  p.  48.) 

It  is  diplacanthid  and  has  several  (two  or  more)  rows  of  inter- 
actinal plates ;  its  dorsal  plates  form  unequal  longitudinal  rows,  each 
plate,  in  the  adults,  bearing  a  transverse  group  of  short,  unequal 
spines,  becoming  numerous  in  large  specimens.  The  minor  pedicel- 
lariae are  abundant  around  the  spines  and  on  the  dermis,  and  there 
are  many  large,  scattered  major  pedicellariae,  some  of  them  unguicu- 
late or  dentate. 


SHALLOW-WATER   STARFISHES  359 

It  varies  much  in  appearance,  according  to  age,  and  has  many  local 
or  casual  varieties.  Whether  all  the  forms  referred  to  it  by  Leipoldt 
and  others  really  belong  together  seems  to  me  somewhat  uncertain. 

I  have  studied  a  series  of  young  and  old  from  Eden  Harbor,  Pata- 
gonia, sent  by  the  Museum  of  Comparative  Zoology.  These  show 
wide  variations,  especially  in  the  character  of  the  dorsal  rows  of 
spines  and  the  number  of  spines  on  each  plate.  The  number  increases 
rapidly  with  age,  so  that  the  larger  specimens  (radius,  175  mm.) 
have  very  numerous,  crowded,  transverse  groups  of  spines  and  pedi- 
cellariae,  the  median  and  some  of  the  others  forming  thick,  wide,  longi- 
tudinal rows.  The  rnadreporic  plate  is  surrounded  by  a  circle  of  small 
special  spines,  increasing  in  number  with  the  age,  from  six  or  seven 
up  to  twenty  or  more. 

The  major  pedicellariae  are  large  and  stout.  Some  are  obtusely 
lanceolate  or  ovate,  others  are  blunt  and  unguiculate,  with  few  teeth. 
They  are  abundant  on  the  edge  of  the  grooves,  where  many  have  long, 
slender  pedicels,  and  between  the  ventral  and  lateral  spines,  and  on 
the  dorsal  papular  areas,  where  they  are  often  stouter  than  the  adja- 
cent spines.  Minor  pedicellariae  are  abundant  on  the  dorsal  dermis, 
crowded,  especially  over  the  transverse  ossicles  and  between  the 
spines. 

The  larger  specimens  may  have  six  to  eight  interactinal  spines  in  a 
transverse  row,  arising,  apparently,  from  three,  to  five,  rows  of  plates. 
Young  specimens,  with  the  greater  radius  50  mm.,  have  only  two 
rows  of  interactinal  plates,  each  with  one  or  two  spines,  and  the 
spines  on  both  marginal  rows  stand  partly  singly,  partly  two  to  a 
plate ;  there  are  but  three  dorsal  rows,  and  the  spines  in  these  stand 
partly  in  pairs,  partly  singly. 

This  species  occurs  on  both  coasts  of  Patagonia  and  Fuegia,  from 
low  tide  apparently  to  348  fathoms. 

COSMASTERIAS  TOMIDATA  (Sladen). 
Cosmasterias  tomidata  SLADEN,  1889,  p.  576. 

This  has  more  regular  rows  of  small,  clustered  dorsal  spines,  and 
very  numerous  dermal  minor  pedicellariae,  larger  than  usual.  The 
major  pedicellariae  are  numerous,  large  and  stout,  often  much  larger 
than  the  spines ;  mostly  strongly  felipedal. 

Gulf  of  Penas,  45  fathoms,  Sladen ;  Porto  Lagunas,  etc.,  50  to  80 
meters,  Leipoldt ;  off  Argentina,  S.  lat.  44°  52',  55  fathoms,  coll.  Mus. 
Comp.  Zoology. 


3OO  VERRILL 

COSMASTERIAS  POLYGRAMMUS   (Sladen,  1889)  Verrill. 
Stichaster  polygrammus  SLADEN,  op.  cit.,  1889,  p.  434,  pis.  c,  cm. 

A  diplacanthid  species,  easily  distinguished  by  the  conspicuous 
transverse  or  banded  arrangement  of  the  dorsal  and  lateral  spines  and 
plates.  The  dorsals  and  superomarginals  form  about  seven  longi- 
tudinal rows.  The  spines  are  small  and  numerous,  arranged  in  nar- 
row transverse  rows  of  three  to  seven  on  the  ossicles.  The  major 
pedicellariae  are  large;  some  of  them  are  felipedal.  Papular  areas 
are  in  regular  rows.  Ventral  plates  bear  four  or  five  spines  in  the 
transverse  rows.  Position  of  genital  pores  unknown. 

The  type  was  from  245  fathoms,  west  of  Magellan  Strait. 

Leipoldt  considered  this  a  form  of  Cosmasterias  sulcifera.  The 
differences  are  notable. 

COSMASTERIAS  BRANDTI  (Bell)  Verrill. 
Cosmasterias  brandti  (BELL,  1881,  p.  91,  pi.  ix,  fig.  i,  as  Asterias). 

The  dorsal  and  lateral  plates  are  said  to  be  about  eleven  in  each 
transverse  row,  and  to  bear  small  spines,  in  transverse  series.  It  may 
be  the  same  as  C.  polygrammus. 

Trinidad  Channel,  at  30  fathoms. 

COSMASTERIAS  FERNANDENSIS  Meissner. 

Cosmasterias  fernandensis  MEISSNER,  1896,  p.  104,  pi.  vi,  fig.  I,  as  Asterias. 
[Polyasterias]  fernandensis  de  Lx>riol,  op.  cit,  1904,  p.  41,  pi.  in,  figs. 
4-8  (rays  unequal,  five  or  six). 

This  has  been  recorded  by  Loriol  from  Gulf  San  Mathias,  E.  Pata- 
gonia. The  type  was  from  Juan  Fernandez.  It  appears  to  be  auto- 
tomous,  though  often  five-rayed. 

GASTRASTER  STUDERI  de  Loriol. 
Gastraster  studeri  DE  LORIOL,  1904,  p.  34,  pi.  iv,  figs.  3-3/. 

This  has  four  irregular  rows  of  podia  proximally,  but  only  two 
rows  on  the  distal  half  of  the  rays.  Its  relations  are  obscure.  It  may 
be  allied  to  Granaster  biseriatus  Krehler. 

From  Gulf  San  Mathias. 

Genus  Adelasterias  Verrill,  nov. 
Type,  A.  papillosa  (Koshler,  1905,  as  Diplosterias) . 
This  is  diplacanthid  and  is  covered  everywhere  with  a  thick  der- 
mis,  which  rises  into  numerous  papilliform  processes,  both  on  the 


t^Lc^C^^ti    /  £r£_ci,*^*>SiMA  )  .  '   J-H--9  *     l; /t.'<<  7 


SHALLOW- WATER   STARFISHES  361 

back  and  on  the  sides,  which  enclose  rudimentary  or  nearly  abortive 
spinules.    The  skeleton  is  almost  entirely  abortive ;  marginal  plates 
are  not  present,  unless  as  rudiments. 
The  type  is  from  the  Antarctic  Ocean. 

Genus  Podasterias  Perrier,  1896. 

Type,  P.  lutkeni  Perrier. 
Diplasterias  (pars)  PERRIER,  1891,  p.  K,  77.    Ludwig,  Voy.  Belgica,  1903. 

This  genus  is  here  extended  to  include  several  paedophoric  dipla- 
canthid  starfishes  having  a  reticulated  dorsal  skeleton,  not  concealed 
by  a  thick  dermis,  and  usually  two  or  more  rows  of  interactinal  plates 
and  spines.  Thus  they  have  a  general  resemblance  to  typical  Asterias, 
like  A.  rubens.  As  some  of  the  species  are  known  to  carry  their 
young,  the  genital  pores  are  presumed  to  be  ventral,  but  I  do  not 
know  that  any  species  has  been  dissected  with  reference  to  this. 
Among  the  species  that  appear  to  belong  here  are  the  following : 

,-P.  lutkeni  (Perrier,  1891,  p.  K,  81,  as  Diplasterias). 

Fuegian  and  Falkland  Is.,  95  to  220  meters. 

Podasterias  steineni  (Studer,  1885,  pi.  i,  figs.  40,  b,  as  Asterias).  Diplasterias 
steineni  Perrier,  1891,  p.  K,  84. 

South  of  Cape  Horn,  99  meters  (Perrier). 
P.  loveni  (Perrier,  1891,  p.  K,  80,  as  Diplasterias). 

S.  lat.  52°  44'  31",  320  meters. 
P.  spinosa  (Perrier,  1891,  p.  K,  82,  as  Diplasterias). 

S.  lat.  47°  29',  depth  not  recorded. 
P.  meridionalis  (Perrier,  1875,  p.  340,  as  Asterias).   7Cu«v^4tJV. 

Antarctic  Expedition. 

Some  of  the  several  species,  from  the  same  region,  described  by 
Bell  in  1881,  probably  also  belong  here,  but  the  skeletal  structure  was 
not  described. 

The  first  two  species  have  been  found  carrying  their  young. 

The  name  Diplasterias  Perrier  is  not  adopted  for  this  genus  because, 
as  originally  defined,  its  only  character  was  the  existence  of  at  least 
two  rows  of  adambulacral  spines.  This  made  it  synonymous  with 
typical  Asterias,  which  was  the  evident  intention  of  Perrier,  for  he 
restricted  Asterias  to  the  monacanthid  species. 

In  the  second  place,  his  first  species  was  A.  sulcifera,  which  he 
evidently  regarded  as  the  type.  But  Sladen's  report  was  published 
while  his  was  being  put  in  type.  So  that  in  an  Appendix,  p.  K,  160, 
he  admits  the  priority  of  Cosmasterias  for  sulcifera  and  abandons 
both  Diplasterias  and  Podasterias  (MS.  name)  and  refers  lutkeni 
(non  Stimpson)  and  its  congeners  to  Asterias.  This  will  not  hold 


362  VERRILL 

good,  for  his  liitkenii  is  paedophoric.  In  a  later  work  (1896,  p.  35)  he 
revived  Podasterias,  with  P.  lutkeni  Perrier  as  the  type. 

Fisher,  1908,  p.  89,  considered  liitkeni  Perrier  a  Pisaster,  and  sug- 
gested a  change  of  name,  owing  to  the  priority  of  Pisaster  liitkenii 
(Stimpson).  They  are  not  congeneric,  the  latter  being  monacanthid, 
and  therefore  no  change  is  needed  in  the  specific  name. 

Genus  Cryptasterias  Verrill,  nov. 
Type,  Diplasterias  turqueti  Koehler,  1905,  p.  465. 

This  is  diplacanthid  and  has  a  reticulated  dorsal  skeleton  entirely 
concealed  by  a  thick  dermis.  The  dorsal  surface  bears  soft  dermal 
papillae,  some  of  which  bear  a  few  pedicellariae ;  others,  scattered,  con- 
tain rudimentary  spinules.  There  is  a  simple  row  of  superomarginal 
spines  and  a  double  row  of  inferomarginals ;  apparently  there  are 
no  interactinal  plates.  It  is  supposed  to  incubate  its  young.  Antarctic. 

STICHASTER  STRIATUS  Miiller  and  Troschel,  1840. 

Asterias  aurantiaca  MEYEN,  Reise  um  die  Erde,  i,  p.  292,  1834  (non  Linne). 
Stichaster  striatus  MULLER  and  TROSCHEL,  op.  cit,  1840,  p.  321  (non  Asterias 

striatus  Lam.). 

Stichaster  aurantiacus  VERRILL,  1867,  p.  293.    Clark,  1910,  p.  337,  pi.  vm,  fig.  i. 
Tonia  atlantica  GRAY,  op.  cit.,  1840,  p.  180;  Synopsis,  1866,  p.  2. 

This  is,  perhaps,  the  most  abundant  littoral  starfish  found  on  the 
entire  coast  of  Chile,  south  to  Talcahuano,  extending  northward  to 
Callao,  Peru  (Yale  Museum,  F.  H.  Bradley).  Common  at  Valpa- 
raiso, on  rocks.  It  grows  to  large  size,  up  to  12  inches  in  diameter  or 
more. 

It  is  the  true  type  of  the  genus  Stichaster,  being  the  only  species 
named  when  that  genus  was  proposed  in  1840.  Perrier  ( 1894,  p.  131 ; 
1896,  p.  27)  -was  in  error  in  making  Asterias  rosea  the  type.  (See 
also  Introduction,  p.  40.) 

It  is  not  generically  related  to  any  North  Pacific  species.  It  is  per- 
haps the  only  known  species  that  can  be  properly  referred  to  the 
genus  Stichaster.  Nearly  all  those  referred  to  it  by  Sladen  and  others 
have  already  been  separated  generically,1  except,  perhaps,  S.  polyplax 
of  the  Australian  region.  This  should  clearly  be  generically  distinct. 
I  would  propose  for  it  the  name  Allo Stichaster. 

*  Perrier,  1894,  proposed  the  genus  Granaster  for  Stichaster  nutrix  Std. ;  but 
G.  biseriatus  Koehler  (antarctic)  seems  to  be  a  distinct  genus,  as  it  has  only 
two  rows  of  podia.  It  may  be  called  HEMIASTERIAS  with  biseriatus  as 
the  type.  •..'» 


SHALLOW-WATER   STARFISHES  363 

Genus  Allostichaster  Verrill,  nov. 
Type,  A.  polyplax  Miiller  and  Troschel. 

It  is  diplacanthid  and  multiple-rayed,  with  one  to  five  madreporic 
plates,  and  is  probably  autotomous.  The  two  rows  of  marginal  plates 
are  stout  and  imbricated ;  dorsal  plates  and  spines  form  five  somewhat 
irregular  longitudinal  rows,  several  short,  obtuse  spines  on  each  plate. 
Minor  pedicellariae  are  dermal,  usually  not  circumspinal. 

SPINULOSA. 

The  Echinasteridae  are  well  represented  by  Henricia,  several 
species ;  Poraniopsis,  one  species ;  on  the  coast  of  Chile,  by  Echinas- 
ter;  at  the  Falkland  Islands,  by  Cribraster  sladeni  Perrier. 

Henricia  seems  to  have  several  species  and  subspecies,  and  these 
vary  much  as  do  the  northern  species. 

H.  obesa  (Sladen,  1889)  was  described  as  from  a  depth  of  from  12 
to  245  fathoms.  Magellan  Strait ;  Falkland  Islands. 

H.  pagenstecheri  (Studer)  was  originally  from  South  Georgia. 
Shallow  water  (  ?) . 

•H.  hyadesi  (Perrier  1891,  as  Cribrella),  Clark,  1910,  p.  337,  pi.  n, 
fig.  5.  This  was  recorded  by  Perrier  from  Fuegia,  etc.,  and  by  Clark 
from  Chile  (ex.  Meissner)  ;  35  to  200  meters  (Perrier). 

H.  studeri  (Perrier,  1891),  from  Magellan  Straits,  in  9  to  183 
meters. 

Ludwig,  1905,  seems  to  consider  all  these  as  variations  of  H.  pagen- 
stacheri.  To  me  most  of  them  seem  as  distinct,  if  correctly  described 
and  figured,  as  some  of  them  are  from  the  corresponding  Arctic 
forms.  These  species  of  Henricia  are  all  pretty  closely  related  to  the 
northern  and  Arctic  species,  contrary  to  those  of  the  Asteriidse. 

Cribraster  is  known  only  from  the  Falkland  Islands;  depth  not 
known,  probably  littoral.  C.  sladeni  Perrier,  1901  (p.  K,  104,  pi.  xi, 
figs.  2a,  zb,  and  Appendix,  p.  K,  161). 

Echinaster  antonioensis  Lor.,  1904,  and  E.  lepidus  Lor.,  1904,  have 
been  described  from  th£  Gulf  of  San  Mathias.  E.  lepidus  is  said  to 
lack  interradial  papular  pores,  and  in  that  case  it  belongs  to  Rhopia 
Gray,  near  R.  sepositus,  of  the  Mediterranean.  The  former  species 
is  related  to  E.  brasiliensis. 

Poraniopsis  echinaster  Perrier,  1891  (p.  K,  106,  pi.  x,  figs.  2,  2a, 
95  meters).  This  is  nearly  related  to  P.  inflata  Fisher,  from  the 
North  Pacific. 

Poraniopsis  mira  (=Lahillia  mira  de  Loriol,  op.  cit,  1904,  p.  33, 
pi.  in,  figs.  i-ih). 


364  VERRILL 

This  is  a  large  species  with  short,  stout  rays,  from  the  Gulf  San 
Mathias,  E.  Patagonia. 

Perknaster  densus  Sladen.    Off  Kerguelen  Island;  127  fathoms. 
P.  fuscus  Sladen.    Off  Heard  Island  and  Kerguelen  Island ;  25  to 
75  fathoms. 

The  family  Acanthasteridae,  pertaining  almost  entirely  to  tropical 
seas,  and  always  rather  rare,  is  said  to  be  represented  by  a  species  in 
the  Magellan  Straits,  but  the  only  record  seems  to  be  an  ancient  one 
and  may  not  be  reliable.  It  has  not  been  found  there  by  any  recent 
expedition. 

Acanthaster  Solaris  (Schr.)  Duj.  et  Hupe  (pars)  was  described  by 
Gray,  1866  (as  Echinaster  Solaris),  from  this  region,  but  apparently 
without  any  modern  record.  According  to  him,  it  has  twenty-one 
rays  and  ten  madreporic  plates.  Duj.  and  Hupe  united  all  the  known 
species  under  this  name. 

A.  ellisii  (Gray)  occurs  in  the  Panamic  fauna.  It  usually  has  from 
eleven  to  thirteen  rays. 

The  oldest  known  species,  A.  planci  (Linne,  Syst.  Nat.,  ed.  X, 
Appendix,  p.  823,  as  Asterias  planci),  occurs  apparently  throughout 
most  of  the  Indo-Pacific  Oceans,  in  the  tropical  zone.  The  type  was 
described  by  Linne  as  having  fifteen  rays.  It  came  from  Goa,  Por- 
tuguese India,  and  the  figure  of  Columna  (Phytobasanos,  pi.  xxxm, 
fig.  A)  was  referred  to  as  its  basis.  The  figure  is  characteristic  and 
the  name  should  be  adopted.  It  is  the  same  as  A.  echinus  (Ellis 
and  So\.)=A.  echinites  of  Lam.  and  of  many  recent  writers. 

The  number  of  rays  and  madreporic  plates  is  variable.  Usually 
there  are  from  fifteen  to  twenty  very  spinose  rays. 

The  family  Asterinidae  is  unusually  well  represented.  On  the 
coast  of  Peru  and  southward  Patiria  chilensis  (Liitken)  and  P.  gayi 
(Perrier)  are  recorded.  They  are  closely  related  and  probably  not 
dinstinct  species;  also,  Patiriella  calcarata  (Perrier)  and  P.  pusilla 
(Perrier).  Desmopatiria  flexilis  Verrill  probably  occurs  on  the 
same  coast,  but  its  origin  is  not  positively  known. 

Farther  south,  on  the  coasts  of  Chile  and  Magellan  Strait,  and 
around  Tierra  del  Fuego  to  Cape  Horn,  the  very  common,  small,  lit- 
toral and  shallow-water  species  is  Patiriella  fimbriata  (Perrier)  or 
P.  bispinosa  (Perrier),  if  the  latter,  a  supposed  variety,  proves  to 
be  distinct. 

P.  fimbriata,  as  originally  described,  was  labelled  as  from  Bourbon 
Island,  but  Perrier  himself  expressed  doubt  as  to  whether  the  locality 
labels  were  correct,  owing  to  the  close  agreement  with  specimens 
from  Chile. 


SHALLOW-WATER  STARFISHES  365 

The  type  had  mostly  a  single  spine  on  the  ventral  plates,  while  the 
Patagonian  form  usually  has  two  spines  on  many  of  the  plates.  There 
are,  also,  some  differences  in  the  dorsal  spinulation,  so  that  they  may 
prove  to  be  distinct;  yet  I  am  not  aware  that  it  has  been  found  in 
tropical  seas  by  later  collectors,  and  the  ancient  Bourbon  Island  label 
may  have  been  wrong.  I  have  examined  numerous  specimens  from 
Magellan  Strait,  and  found  but  little  variation,  but  have  seen  no 
authentic  tropical  specimens.  It  has  a  thicker  dermis  than  usual,  and 
is  not  a  typical  Patiriella. 

P.  calcarata  and  P.  pusilla  are,  however,  more  nearly  allied  to  sev- 
eral African  and  Australian  species  than  to  any  in  the  northern  hemi- 
sphere. 

Asterinides  perrieri=Asterina  perrieri  de  Loriol,  1904,  a  small 
species  from  E.  Patagonia,  Gulf  of  San  Mathias,  appears  to  belong 
to  the  genus  Asterinides,  but  may  be  closely  related  to  P.  fimbriata 
or  bispinosa. 

Enoplopatiria  marginata,  which  has  been  recorded  from  near  the 
eastern  end  of  Magellan  Strait,  is  a  common  Brazilian  species,  extend- 
ing northward  to  the  West  Indies,  and  recorded  also  from  the  west 
coast  of  Africa.  A  closely  related  new  species,  E.  siderea,  in  our  col- 
lection, is  labeled  as  from  Panama  (donor,  Captain  Dow,  of  the 
Panama  and  California  steamship  line). 

E.  siderea  Verrill  is  a  rather  large  species,  radii,  24  mm.  and  48  mm., 
with  stout,  short,  broad  rays  and  well  defined  papular  areas,  with 
numerous  forked  dorsal  pedicellariae ;  ventral  plates  with  a  comb  of 
three  elongated  spines ;  four  webbed  furrow  spines. 

It  differs  from  E.  marginata  in  having  the  spinules  of  the  inter- 
radial  dorsal  plates  longer,  widely  divergent,  stellate  and  paxilliform, 
and  in  other  characteristics ;  but  it  is  evidently  closely  related,  so  much 
so  that  I  formerly  suspected  the  correctness  of  the  label,  thinking  that 
it  might  have  come  from  Colon,  on  the  Atlantic  side,  and  might  be 
only  a  large  variety  of  E.  marginata.  See  plate  cix,  figure  4. 

The  family  Ganeriidae  is  very  characteristic  of  the  Patagonian- 
Fuegian  fauna.  Indeed,  nearly  all  the  known  genera  and  species 
are  from  that  region  and  adjacent  districts. 

Of  the  genus  Ganeria  Gray,  1847,  Perrier,  1891,  recognized  four 
species;  viz.:  G.  falklandica  Gray,  1847,  Falkland  Islands;  east  of 
Magellan  Strait,  55  fathoms,  Sladen;  G.  robusta  Perrier,  1891,  28 
meters;  G.  hahni  Perrier,  1891,  138  meters;  G.  papillosa Perrier,  1891, 
depth  not  given. 


366  VERRILL 

The  genus  Lebrunaster  Perrier,  1891,  p.  K,  116,  type  L.  paxillosus 
Perrier,  from  Patagonia,  depth  unknown,  is  known  only  from  this 
fauna. 

Numerous  nominal  species  of  Cycethra  have  been  described  from 
Magellan  Strait  and  the  coasts  of  Patagonia  and  Terra  del  Fuego, 
where  the  genus  is  common  and  variable. 

Ludwig,  1905,  reduces  them  mostly  to  varieties  of  C.  verrucosa 
(Phil.).  This  species  includes  C.  nitida  Sladen,  Meissner  -f-  C.  elec- 
tilis  Sladen,  Leipoldt  -+-  C.  simplex  Bell,  Meissner  -f-  C.  pinguis 
Sladen  -f-  C.  ganeriodes  Perrier  -f-  C.  elongata  Perrier  -j-  C.  media 
Perrier -f C.  asterina  Perrier +C.  subelectillis  Perrier +C.  calva  Per- 
rier -f-  C.  regularis  Perrier  -f  C.  asteriscus  Perrier.  The  last  eight 
forms  were  regarded  by  Perrier  himself  as  mere  varieties  of  C.  sim- 
plex, as  well  as  some  of  the  others. 

They  are  common  in  Magellan  Strait  and  southward  to  Cape 
Horn,  from  low  tide  to  40  fathoms.  Cycethra  lahillei  de  Loriol, 
1904,  p.  25,  pi.  II,  fig.  2,  from  Gulf  of  San  Mathias,  appears  to  be 
a  distinct  species,  with  short  rays,  imperfectly  developed  marginal 
plates,  and  slender  paxilliform  spinules. 

One  of  the  generic  types,  common  to  this  region  and  the  North 
Pacific,  is  Peribolaster,  of  the  family  Korethrasteridae,  of  which  only 
two  species  are  known.  P.  foliculata  Sladen  occurred  in  45  fathoms, 
west  of  Patagonia.  P.  biserialis  Fisher,  1911,  ranges  from  Cali- 
fornia to  Bering  Sea,  in  from  57  to  313  fathoms. 

The  family  Solasteridae  is  represented  in  moderate  depths  by 
Solaster  australis  (Perrier)  and  Lophaster  stellans  Sladen,  both  of 
which  have  occurred  in  from  30  to  40  fathoms.  In  greater  depths 
several  additional  species  of  those  genera  occur.  (See  list  of  deeper- 
water  species  below,  p.  368.) 

These  are  genera  widely  distributed  in  all  seas,  especially  in  rather 
deep  water,  coming  into  shallow  water  only  in  the  colder  regions. 
Some  of  the  Antarctic  species  are  closely  allied  to  those  of  the  North 
Pacific  and  Arctic  Ocean,  like  those  of  other  families  from  deep  water. 

The  Pterasteridse  are  also  well  represented  by  several  species  of 
Pteraster  and  Diplopteraster  in  waters  of  somewhat  greater  depths. 
These  are  also  allied  to  the  corresponding  species  of  the  North 
Atlantic  and  North  Pacific.  ( See  list  of  deeper- water  species  below, 
p.  368.) 

The  genus  Retaster,  which  is  represented  by  two  species,  does  not 
occur  in  northern  waters,  but  is  found  in  the  Indo-Pacific  region, 
from  Cape  Good  Hope  to  Australia.  The  Patagonian  species  are  as 
follows : 


SHALLOW-WATER   STARFISHES  367 

R.  gibber  Sladen,  1889;  Ludwig,  1905,  pi.  v,  figs.  6,  7.  This 
occurred  in  from  14  to  245  fathoms,  in  Magellan  Strait. 

R.  verrucvsus  Sladen.  This  occurred  in  55  fathoms,  in  Magellan 
Strait. 

PHANEROZONA. 

The  family  Odontasteridae  is  better  represented  in  the  Fuegian 
fauna  than  in  most  regions,  some  species  occurring  commonly  even  at 
low  tide  and  in  shallow  water.  Several  nominal  species  have  been 
described,  some  of  which  Ludwig  (1905)  and  others  reduce  to  syno- 
nyms. Ludwig  recognizes  three  shallow-water  species ;  viz. : 

1.  O.  grayi  (Bell,  non  Perrier)  =  Gnathaster  grayi  Sladen  =  O. 
pedicettaris  (Perrier). 

2.  O.  penicillatus  (Phil.)  -f-  O.  pilulatus  (Sladen,  as  Gnathaster) 
-\- O.  meridionalis  (Smith).    Leipoldt,  1895. 

3.  Asterodon  singularis  (Miiller  and  Troschel)  Leipoldt,  1895,  pi. 
xxxi,  figs.  7&-c,  as  Odontaster  =  A.  granulosus  Perrier,  1891,  pi.  xi, 
figs.  4a,  b. 

A.  belli  (Studer)  Perrier  is  probably  not  distinct  from  A.  singularis. 

The  first  species  is  from  a  depth  of  10  to  97  meters.  The  second 
species  ranges  from  5  to  150  fathoms.  Several  other  antarctic  species 
have  been  described  from  deeper  water.  ( See  Sladen ;  also  Koehler, 

I905-) 

Luidia  magellancia  Leipoldt,  1895,  p.  610,  pi.  xxxn,  figs.  na-c. 
This  is  remarkable  as  inhabiting  cold  waters,  for  all  the  other  numer- 
ous species  inhabit  tropical  or  temperate  seas.  The  type  had  the 
radii  208  m.  and  35  m ;  ratio,  1 : 6.  Its  nearest  related  species  are  L. 
bellona  Liitken,  which  ranges  from  Chile  to  Mexico,  and  L.  phragma 
Clark  (1910,  p.  329,  pi.  n,  fig.  i),  which  was  supposed  to  be  from 
Peru.  This  may,  therefore,  be  considered  as  one  of  the  few  that  were 
probably  of  Panamic  origin. 

IX.  LIST  OF  PATAGONIAN  AND  FUEGIAN »  STARFISHES  FOUND 
IN  FROM  SO  TO  500  FATHOMS,  MOSTLY  RELATED  TO 
NORTHERN  OR  ARCTIC  SPECIES. 

In  the  somewhat  deep,  cold  waters  on  the  "  continental  slopes  "  (50 
to  500  fathoms),  a  considerable  number  of  genera  and  species  have 
been  found  that  are  closely  related  to  those  of  the  North  Pacific  and 
North  Atlantic  and  the  Arctic  Ocean.  Many  of  these  have  been 
recorded  by  Sladen,  1889;  Perrier,  1891;  Leipoldt,  1895;  Ludwig, 
1905 ;  Koehler,  1905,  and  others.  Among  them  are  the  following : 

1  Some  species  from  Kerguelen  Island,  etc.,  are  here  included,  but  some  are 
common  to  the  two  regions. 


368  VERRILL 

Coscinasterias  candicans  (Lud.,  1905,  as  Stolosterias). 

S.  lat.  70°  to  71°  ;  450  to  560  meters. 
Pedicellaster  antarcticus  Lud.,  1905. 

430  to  450  meters. 
P.  scaber  (Smith)  Perrier. 

Kerguelen  L,  etc. 
Henricia  simplex  (  Sladen,  1889,  as  Cribrella) . 

Antarctic ;  50  to  310  fathoms. 
H.  prcestans  (Sladen,  as  Cribrella). 

210  fathoms. 
H.  hyadesi  (Perrier,  1891). 

35  to  200  fathoms. 
Echinaster  smithii  Ludwig. 

450  meters. 
Solaster  octoradiatus  Ludwig. 

450  to  500  meters;  S.  lat.  70°  to  71°. 
Solaster  subarcuatus  Sladen. 

150  fathoms. 
Solaster  regularis  Sladen. 

175  fathoms. 
5".  australis  (Perrier). 

65  to  198  meters. 
Lophaster  stellans  Sladen. 

40  to  1325  fathoms.    Includes  L.  pentactis  Per.,  200  meters. 
Pteraster  stellifer  Sladen. 

245  fathoms. 
P.  rugatus  Sladen. 

150  fathoms. 
P.  ingouffi  Per. 

Fuegian ;  270  meters. 
P.  lebruni  Per. 

Fuegian;  80  meters;  450  meters,  Ludwig;  S.  lat.  71°  24'. 
Diplopteraster  verrucosus  (Sladen,  as  Retaster). 

East  of  Magellan  Strait ;  55  fathoms. 
D.  peregrinator  (Sladen,  as  Retaster). 

Kerguelen  I. ;  127  fathoms.  , 

Hymenaster  perspicuus  Lud. 

400  to  450  meters. 
Porania  antarctica  Smith. 

Kerguelen  I.;  South  Georgia;  Magellan  Strait;  Antarctic  Ocean,  S.  lat. 
71°  18',  450  meters;  Fuegian,  10  to  1,600  fathoms.    Includes  P.  ma- 
gelhanica  Studer  and  P.  patagonica  Perrier. 
P.  glaber  Sladen. 

30  to  127  fathoms. 
Odontaster  cremens  Ludwig. 

450  meters. 
Mimaster  cognatus  Sladen. 

245  to  1325  fathoms ;  450  meters,  S.  lat.  70°  23',  Ludwig. 
Pseudar 'chaster  patagonicus  (Per.,  1891,  p.  K,  125,  pi.  xm,  figs.  20,  2b,  as 
Astrogonium-) . 

283  meters. 


SHALLOW-WATER   STARFISHES  369 

Hippasteria  hyadesi  Per. 

326  meters. 
Ceramaster  patagonicus  (Sladen). 

55  to  245  fathoms. 
C.  austro-granularis  (Per.,  1891,  as  Pentagonaster). 

340  meters. 
Leptychaster  kerguelenensis  Smith. 

Kerguelen  and  Marion  Islands ;  10  to  100  fathoms. 
L.  antarcticus  Sladen. 

210  fathoms. 
Psilaster  fleuriaisi  (Per.,  1891,  as  Goniopecten). 

198  to  283  meters. 
Cheiraster  gerlachei  Lud. 

450  to  560  meters. 
Pontaster  planeta  Sladen. 

245  fathoms. 
Ctenodiscus  australis  Liitk. 

55  to  600  fathoms. 
C.  procurator  Sladen. 

40  to  1325  fathoms. 

All  the  species  (thirty-four)  included  in  this  list  are  generically 
allied  to  northern  or  arctic  species.  In  many  cases  the  resemblance  is 
very  close  specifically.  This  is  so  especially  in  the  cases  of  some 
species  of  Henricia,  Solaster,  Pteraster,  Diphpteraster,  Porania, 
Hippasteria,  Ceramaster,  and  Ctenodiscus. 

These  facts  clearly  indicate  that  extensive  migrations  of  the  ances- 
tors of  these  species  have  occurred,  in  the  colder  deep  waters,  in  one 
or  both  directions,  at  no  very  remote  geologic  periods. 

It  appears,  with  our  present  knowledge  of  oceanic  currents  and 
temperatures,  that  the  greater  part  of  the  migration  would  have  been 
from  the  Arctic  Ocean  southward  to  the  Antarctic.  But  no  barriers 
exist,  at  the  depths  that  these  species  inhabit,  to  prevent  a  northward 
migration,  for  abyssal  species  are  not  included  in  the  list.  The  shal- 
low-water and  littoral  species,  on  the  other  hand,  show  no  evidence 
of  such  comparatively  recent  migrations,  for  much  the  larger  part 
are  peculiar  to  this  region  and  the  adjacent  Antarctic  seas  and  islands, 
with  a  few  additions  from  the  Indo-Pacific  region. 

Scarcely  any  of  the  strictly  shallow-water  species  seem  to  have 
been  able  to  cross  the  tropical  regions,  on  either  side  of  America, 
unless  in  remote  geologic  time. 

Those  that  can  descend  to  the  depths  where  very  cold  water  exists, 
even  in  tropical  seas,  have  been  able  to  migrate  from  polar  seas  across 
the  tropics,  probably  in  both  directions,  no  doubt  with  extreme  slow- 
ness, and  yet  with  but  little  change  in  appearance  or  structure.  Such 

25 


37°  VERRILL 

migrations  have  probably  been  going  on  during  vast  geologic  periods 
and,  no  doubt,  are  still  in  progress. 

Many  of  the  existing  genera  of  starfishes  are  doubtless  of  great 
antiquity,  geologically  speaking,  and  show  wonderful  persistence  in 
many  of  the  details  of  structure  and  habit. 

X.  LISTS  OF  EXTRALIMITAL  SPECIES  PARTIALLY  DESCRIBED, 
REVISED,  OR  FIGURED. 

Many  other  extralimital  species,  merely  included  in  the  faunal 
lists,  or  only  mentioned  by  name  under  their  genera,  are  not  included 
in  these  lists. 

I.    NORTH  ATLANTIC  AND  ARCTIC  SPECIES. 

Asterias  vulgaris  Ver. 

New  England. 
Asterias  forbesi  Desor. 

New  England,  etc. 
A.  acervata  borealis  =  A.  polaris. 

Arctic  and  North  Atlantic. 
Orthasterias  tanneri  Ver. 

Off  East  coast  United  States.    PI.  XLVIIJ,  fig.  6. 
O.  subangulosa  Ver.  (nom.  nov.)  =A.  angulosa  Per.,  preoccupied. 

West  Indies. 

Stylasterias  neglect  a  (Per.  non  Bell). 
Coscinasterias  tenuispina  (Lam.)  Ver. 

Europe;  Bermuda. 
Marthasterias  glacialis  (Mull.)  Jul. 

Europe,  etc. 
Sclerasterias  guernei  Per. 

East  Atlantic. 
Leptasterias  littoralis  (Stimp.)  Ver. 

New  England. 
L.  compta  (Stimp.)  Ver. 

New  England. 
L.  tenera  (Stimp.)  Ver. 

New  England. 
Ctenasterias  spitzbergensis  (D.  and  Kor.). 

Arctic. 
Urasterias  linckii  (M.  and  Tr.)  Ver. 

Nova  Scotia;  Arctic  Ocean;  post-glacial  fossil  at  Salem,  Mass.    PI.  LXX, 

figs.  1-4. 
U.  panopla  (Stuxb.)  Ver. 

Arctic. 
Stichastrella  rosea  (Mull.)  Ver. 

Europe. 
Coronaster  briareus  Ver. 

West  Atlantic. 


SHALLOW- WATER   STARFISHES  371 

Henricia  sanguinolenta  miliaris  Ver. 

New  England. 
H.  antillarum  (Per.). 

West  Indies. 
H.  sexradiata  (Per.)  Ver. 

West  Indies. 
Cr ossaster  korenii  Ver.  (nom.  nov.). 

Arctic  and  North  Atlantic. 
Asterina  gibbosa  (Penn.)  Gray. 

Europe. 
A.  Pygmcea  Ver.  =  Porania.  (Young). 

New  England. 
Asterinides  folium  (Liitk.)  Ver. 

West  Indies;  Bermuda. 
Enoplopatiria  marginata  (Hupe)  Ver. 

West  Indies ;  Brazil ;  West  Africa,  etc. 
Tremaster  mirabilis  Ver. 

Newfoundland  Banks,  etc. 
Hippasteria  phrygiana  (Parel.)  Ag. 

New  England;  Europe;  Arctic  Ocean.    PI.  XLVII,  fig.  i;  pi.  XLVIH,  figs. 

1-5 ;  pi.  XLIX,  fig.  6. 
Mediaster  bairdii  Ver. 

West  Atlantic.    PI.  n,  fig.  2;  pi.  in,  fig.  2  (type). 
Blakiaster  conicus  Per. 

West  Indies. 
Luidia  clathrata  (Say). 

Southern  coast  United  States ;  Bermuda ;  West  Indies,  etc.  PL  an,  fig.  i. 

2.    PATAGONIAN,  FUEGIAN,  ANTARCTIC,  AND  SOUTH  ATLANTIC  SPECIES. 

Labidiaster  radiosus  Liitk. 

Pedicellaster  antarcticus  Lud. 

Sporasterias  rttgispina  (Stimp.)   Per. 

S.  antarctica  (Liitk.)  Per. 

S.  perrieri  (Smith). 

5".  rupicola  (Ver.). 

Anasterias  minuta  Per. 

A.  lysasteria  (Ver.),  nom.  nov.  =  A.  perrieri  Std.  (non  Per.)- 

A.  belgica  Ludwig. 

A.  studeri  (Perrier). 

A.  tenera  Koehler. 

A.  verrillii  (Bell). 

Pcedasterias  (gen.  nov.)  chirophora  (Lud.)  Ver. 

Calvasterias  asterinoides  Per. 

C.  (?)  stolidota  Sladen. 

M eyenaster  gelatinosus  (Meyen)  Ver. 

Smilasterias  scalprifera  Sladen. 

Cosmasterias  lurida  (Phil.)  Lud.    Includes  C.  sulcifera,  etc. 

C.  tomidata  Sladen. 

C.  polygrammus  (Sladen,  as  Stic  hosier). 


372  VERRILL 

C.  brandti  (Bell,  as  Asterias)  Ver. 

C.  fernandensis  (Meiss.,  as  Polyasterias)  Ver. 

Gastraster  studeri  Koehler. 

Adelasterias  (gen.  nov.)  papillosa  (Koehler,  1905,  as  Asterias). 

Podasterias  liitkeni  Per. 

P.  meridionalis  (Perrier,  as  Asterias). 

P.  spinosa  (Perrier,  as  Diplasterias) . 

P.  steineni  Per. 

Cryptasterias  (gen.  nov.)  turqueti  (Kcehler,  as  Asterias)  Ver.  Antarctic. 

Granaster  nutrix  (Studer)  Per.    S.  Georgia. 

Hemiasterias  biseriatus  (Koehler,  as  Granaster)  Antarctic. 

Stichaster  striatus  (M.  and  Tr.)  =  Asterias  aurantiaca  Meyen  (non  Linne). 

Peru  and  Chile. 

Heliaster  helianthus  (Lam.)  D.  and  H.    Ecuador  to  Chile. 
Henricia  obesa  (Sladen). 
H.  pagenstecheri  (Studer). 
H.  hyadesi  (Per.)  Clark. 
Cribraster  sladeni  Per. 
Echinaster  antonioensis  Koehler. 
E.  (Rhopia)  lepidus  (Koehler). 
Poraniopsis  echinaster  Per. 
P.  mira  (Koehler,  as  Lahillia). 

Acanthaster  Solaris  (Schr.)  D.  and  Hupe.    Panamic,  etc. 
Patiria  chilensis  (Liitk.)  Ver. 
P.  gayi  (Per.)  Ver. 
Patiriella  calcarata  (Per.)  Ver. 
P.  pusilla  (Per.)  Ver. 
P.  Ambriata  (Per.)  Ver. 
P.  bispinosa  (Per.)  Ver. 

Asterinides  perrieri  (Koehler,  as  Asterina)  Ver. 
Desmopatiria  flexilis  Ver. 
Porania  antarctica  Smith. 
Ganeria  falklandica  Gray. 
G.  papillosa  Per. 
Lebrunaster  paxillosus  Per. 

Cycethra  verrucosa  (Phil.).   Includes  C.  simplex  Bell;  C.  electilis  Sladen,  etc. 
C.  lahillei  Koehler. 
Peribolaster  foliculata  Sladen. 
Solaster  australis  (Per.) 
Lophaster  stellans  Sladen. 
Retaster  gibber  Sladen. 
R.  verrucosus  Sladen. 
Odontaster  grayi  (Bell). 

O.  penicillatus  (Phil.).    Includes  O.  pilulatus  Sladen. 
Asterodon  singularis  (M.  and  Tr.).    Includes  O.  granulosus  Per. 
Leptychaster  kerguelenensis  Smith. 
Luidia  magellanica  Leipoldt. 
Astropectenides  (gen.  nov.)  mesacutus  (Sladen).    South  Atlantic. 


SHALLOW-WATER  STARFISHES  373 

3.    SPECIES  FROM  THE  PANAMIC  FAUNA. 

Coscinasterias  alexandri  (Ludwig,  as  Stolasterias) . 
Stylasterias  robusta  (Ludwig,  as  Stolasterias). 
1  Acanthaster  ellisii  (Gray). 
Mithrodia  bradleyi  Ver.    PI.  cvm,  fig.  i  (type). 
Enoplopatiria  siderea  Ver.    PI.  ax,  fig.  4. 
Echinaster  tenuispinus  Ver.    PI.  cvn,  fig.  2,  type. 
Asterinides  modesta  Ver. 
Callopatiria  obtusa  (Gray)  Ver. 
Amphiaster  insignis  Ver.    PI.  cvin,  fig.  i  (type) . 
Astropecten  oerstedii  Gray. 
A.  armatus  Gray. 

4.   INDO-PACIFIC,  AUSTRALIAN,  AND  JAPANESE  SPECIES. 
Labidastrella  (gen.  nov.)  annulata  (Sladen,  as  Labidiaster) . 

Australasian;  off  Heard  Island  and  Kerguelen  Island,  in  75-150  fathoms.        „  **v~ 

Heterasterias  (gen.  nov.)  volsellata  (Sladen,  as  Asterias). 

Japan. 
Coscinasterias  acutispina  (Stimp.). 

Japan. 
C.  muricata  Ver. 

New  Zealand. 
Allostichaster  (gen.  nov.)  polyplax  (M.  and  Tr.)  Ver. 

Australasian. 
Distolasterias  stichantha  (Sladen)  Ver. 

Japan. 
.  Allasterias  forficulosa  Ver. 

Japan.    PI.  LXXXIII,  figs.  3~3c ;  pi.  LXXXIV,  fig.  I. 
^A.  versicolor  (Sladen)  Ver. 

Japan. 
v  A.  amurensis  (Liitk.)  Ver. 

Siberia. 
A.  migrata  (Sladen,  1878,  as  Asteracanthion  rubens  var.). 

Straits  of  Korea. 
Ccelasterias  australis  Ver. 

New  Zealand. 
Acanthaster  planci  (Linne)=/4.  echinites  of  authors. 

Indo-Pacific. 
Henricides  (gen.  nov.)  heteractis  (Clark,  as  Henricia). 

Lord  Howe  I.;  Australasian. 
Henricia  densispina  (Sladen,  1878). 

Straits  of  Corea ;  40  fathoms. 
H.  japonica  Ver,, 

Japan. 
Patina  coccinea  Gray. 

Indo-Pacific. 
Asterinopsis  penicillaris  (Lam.)  Ver. 

Indo-Pacific ;  Australasian. 
Patiriella  regularis  Ver. 

New  Zealand. 


s)r 


374  VERRILL 

XI.  LIST  OF  NEW  GENERA  ESTABLISHED  AND  DESCRIBED 

Brisingidae  : 

1      Labidastrella.    Type  Labidiaster  annulata  Sladen.     =-  / 

A  steriidae  : 

Evasterias.    Type,  E,  troschelii  (Stimpson). 

Orthasterias.    Type,  O.  columbiana  Verrill. 

Stylasterias.    Type,  O.  /orrm  (de  Loriol).     r*  < 

Heterasterias.    Type,  H.  volsellata  (Sladen). 

Stenasterias.    Type,  S.  macropora  Ver. 
?Y  j  Ctenasterias.    Type,  C.  spitsbergensis  (Danielssen  and  Koren).- 

Tosiaster.    Type,  7\  arcticus  Ver. 

Allostichaster.   Type,  A.  polyplax  (Miiller  and  Troschel). 
.Padasterias.    Type,  P.  chirophora  (Ludwig). 

Adelasterias.    Type,  ^4.  papillosa  (Kcehler). 

Cryptasterias.    Type,  C.  turqueti  (Kcehler). 

Parasterias.    Type,  P.  albertensis  Verrill. 

Stichastrella.    Type,  5.  ro^eo  (Muller). 

*t&'Hemiasterias.    Type,  H.  biseriatus  (Kcehler,  as  Granaster). 
Echinasteridae  : 

Henricides.    Type,  H.  heteractis  (Clark). 
Astropectinidae  : 

Astropectinides.    Type,  A.  mesacutus  (Sladen). 

BIBLIOGRAPHY  OF  STARFISHES  OF  NORTHWEST 
COAST,  AND  OTHER  WORKS  REFERRED  TO. 

As  a  rule,  general  works  and  text-books  are  not  included  in  this 
list,  unless  specially  referred  to  in  the  text. 

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26 


VERRILL 

No.  2.  The  Echinoderms  of  Panama  and  the  West  Coast  of  America, 
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No.  3.  On  the  Geographical  Distribution  of  the  Echinoderms  of  the 
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No.  5.  Notice  of  a  Collection  of  Echinoderms  from  La  Paz,  Lower 
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No.  8.  Additional  Observations  on  Echinoderms  chiefly  from  the  Pa- 
cific Coast  of  America,  pp.  568-593,  pi.  x,  18710. 

No.  9.    The  Echinoderm  Fauna  of  the  Gulf  of  California  and  Cape  St. 
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Geographical  Distribution  of  fifty  species. 

— .  Marine  Fauna  of  Eastport,  Me.  Bull.  Essex  Inst.,  Vol.  in,  pp.  2-6, 
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— .  Results  of  Recent  Dredging  Operations  on  the  Coast  of  New  Eng- 
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— .  Report  upon  the  Invertebrate  Animals  of  Vineyard  Sound,  etc., 
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— .  Note  on  some  of  the  Starfishes  of  the  New  England  Coast.  In 
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Corrections  of  Errors  made  by  M.  E.  Perrier,  in  respect  to  New  Eng- 
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— .  Notice  of  Recent  Additions  to  the  Marine  Fauna  of  the  Eastern 
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— .  Radiates.  In  Ludwig  Kumlien,  Contributions  to  the  Natural  History 
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— .  Preliminary  Check  List  of  the  Marine  Invertebrata  of  the  Atlantic 
Coast,  from  Cape  Cod  to  the  Gulf  of  St.  Lawrence,  32  pp,  1879  (with  two 
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— .  Notice  of  Recent  Additions  to  the  Marine  Invertebrata  of  the  North- 
eastern Coast  of  America,  with  Descriptions  of  New  Genera  and  Species 
and  Critical  Remarks  on  Others.  Proc.  U.  S.  Nat  Mus.,  vol.  11  (1879), 
pp.  165-205,  Washington,  18800. 

— .    List  of  Marine  Invertebrata  from  the  New  England  Coast.     Proc. 
U.  S.  Nat.  Mus.,  vol.  n  (1879),  pp.  227-232,  Washington,  18806. 
See  also  under  Whiteaves,  J.  F.,  i88oc. 

— .  Notice  of  the  Remarkable  Marine  Fauna  Occupying  the  Outer  Banks 
off  the  Southern  Coast  of  New  England,  no.  i,  no.  3,  and  no.  4.  Amer. 
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142 ;  no.  4,  216-225,  New  Haven,  1882. 


SHALLOW-WATER   STARFISHES  387 

— .    The  same,  no.  9,  vol.  xxvm,  pp.  213-220,  18840. 
— .    The  same,  no.  10,  vol.  xxvin,  pp.  378-484,  18846. 

— .    The  same,  no.  n,  vol.  xxix,  pp.  149-157,  18850. 

.     Notice  of  the  Remarkable  Marine  Fauna  Occupying  the  Outer  Banks 

off  the  Southern  Coast  of  New  England,  and  of  Some  Additions  to  the 
Fauna  of  Vineyard  Sound.  In  the  Annual  Report  of  the  U.  S.  Com- 
missioner of  Fish  and  Fisheries  for  1882,  26  pp.,  i884c. 

.    Results  of  the  Explorations  made  by  the  steamer  Albatross  off  the 

Northern  Coast  of  the  United  States  in  1883.  In  the  Annual  Report  of 
the  Commissioner  of  Fish  and  Fisheries  for  1883,  pp.  1-107  (503-699),  44 
pis.,  Washington,  18856. 

Geographical  and  bathymetrical  distribution. 

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Revision  of  Certain  Species  formerly  Described.  Proc.  U.  S.  Nat.  Mus. 
vol.  xvn,  pp.  245-297,  Washington,  1804. 

— .  Distribution  of  the  Echinoderms'of  Northeastern  America.  Amer. 
Journ.  Science,  vol.  XLIX,  pp.  127-141,  199-212,  New  Haven,  18950,  Ab- 
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— .  Revision  of  Certain  Genera  and  Species  of  Starfishes,  with  De- 
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vol.  x,  part  i,  pp.  145-234,  pis.  xxivo-xxx,  New  Haven,  1899. 

— .    Additions  to  the  Fauna  of  the  Bermudas  from  the  Yale  Expedition 
of  1901  (47  pp.,  pis.  i-ix;  6  cuts  in  text),  1901.    Trans.  Conn.  Acad.  Sci., 
vol.  xi,  pp.  15-62,  1901. 
Habits  of  Luidia  clathrata  on  p.  36;  rapid  progression,  etc. 

— .    Zoology  of  the  Bermudas,  vol.  I,  427  pp.,  45  pis.,  1903. 
Starfishes  are  discussed  in  article  10,  p.  36. 

— .    The  Bermuda  Islands,  part  5,  section  i,  Characteristic  Life  of  the 
Bermuda  Coral  Reefs.     Trans.  Conn.  Acad.   Sci.,  vol.  xn,  pp.   160-316, 
28  pis.,  i  map,  120  text  cuts,  1906. 
Starfishes  are  discussed  on  pp.  280-281,  pis.  xxiv,  xxxrwr,  xxxvic. 

— .  Descriptions  of  New  Genera  and  Species  of  Starfishes  from  the 
North  Pacific  Coast  of  America.  (Brief  Contributions  from  the  Museum 
of  Yale  University,  no.  LXX.)  Amer.  Journ.  Sci.,  vol.  xxvin,  pp.  59-70, 
figs,  i-io,  July,  19090. 

Remarkable  Development  of  Starfishes  on  the  Northwest  American 


Coast;  Hybridism;  Multiplicity  of  Rays;  Teratology;  Problems  in  Evo- 
lution; Geographical  Distribution.  The  American  Naturalist,  vol.  XLIII, 
PP-  542-555,  figs.  i-7,  September,  19096. 

.    Revision  of  the  Genera  of  Starfishes  of  the  Subfamily  Asterininae. 

Amer.  Journ.  Science,  ser.  4,  vol.  xxxv,  pp.  477-485,  2  text  cuts,  May,  1913. 
Eight  new  genera  established. 

VIGUIER,  C.  Anatomic  comparee  du  Squelette  des  Stellerides.  Arch.  Zool. 
Exper.  et  Gener.,  vol.  vn,  pp.  33-250,  pis.  v-xvi,  1879. 

WAGNER,  NICOLAS.  Die  Wirbellosen  des  Weifen  Meeres,  Bd.  i,  Leipzig,  1885, 
folio,  171  pp.,  21  pis.,  Enthalt,  pp.  170-171 :  Th.  Jarzynsky,  Catalogus 
Echinodermatum  inventorum  in  man  albo  et  in  mari  glaciali  ad  litus 
murmanicum,  anno  1869  et  1870.  1885. 

-^  WHITEAVES,  J.  F.  Report  on  a  Deep-sea  Dredging  Expedition  [in  1871] 
to  the  Gulf  of  St.  Lawrence.  In  Report  of  Minister  of  Marine  and 
Fisheries  for  Canada,  18720. 


388  VERRILL 


— .  Notes  on  a  Deep-sea  Dredging  Expedition  [in  1871]  round  the  Island 
of  Anticosti  in  the  Gulf  of  St.  Lawrence.  Ann.  Mag.  Nat.  Hist.  (4), 
vol.  x,  pp.  341-354,  18726. 

— .  The  same.  Report  on  a  Second  Deep-sea  Dredging  Expedition  [in 
1872],  22  pp.,  1873. 

— .    The  same.    Third  Report,  29  pp.,  18740. 

— .  On  Recent  Deep-sea  Dredging  Operations  in  the  Gulf  of  St.  Law- 
rence. In  Amer.  Journ.  Science  and  Arts  (3),  vol.  vin,  pp.  210-219,  New 
Haven,  18746. 

— .  On  Some  Marine  Invertebrata  from  the  West  Coast  of  North 
America.  In  Canadian  Naturalist,  vol.  vm,  9  pp.,  December,  1878. 

— .    On  Some  Marine  Invertebrata  from  the  Queen  Charlotte  Islands. 
In  Report  of  Progress  of  Geological  Survey  of  Canada,  1878-79,  16  pp., 
Ottawa,  1880. 
Includes  descriptions  of  three  new  starfishes,  by  A.  E.  Verrill. 

— .  On  Some  Marine  Invertebrata,  Dredged  or  otherwise  Collected  by 
Dr.  G.  M.  Dawson,  in  1885,  on  the  Coast  of  British  Columbia,  etc.  In 
Trans.  Royal  Society  of  Canada,  vol.  iv,  section  iv,  pp.  111-137,  1887. 

Catalogue   of   the   Marine   Invertebrata   of   Eastern    Canada.     In 


Geological  Survey  of  Canada,  272  pp.,  Ottawa,  1901. 
'  XANTUS,  J.     Descriptions  of  Three  New  Species  of  Starfishes  from  Cape 
St.  Lucas.    In  Proc.  Acad.  Nat.  Sciences  Philadelphia,  p.  568,  1860. 
These  are  Asterias  sertulifera,  Heliaster  microbrachia  and  H.  kubiniji. 


INDEX. 


(  Synonyms  in  italic  type.    Descriptive  page  references  in  black  face  type.) 


Acantharchaster  311. 
Acantharchaster  dawsoni  312. 
Acanthaster  207. 
Acanthaster  echinites  364,  373. 
Acanthaster  echinus  364. 
Acanthaster  ellisii  364,  373. 
Acanthaster  planci  364,  373. 
Acanthaster  Solaris  364,  372. 
Acanthasteridae  204. 
acanthostoma,  Evasterias,  new  species 

11,  64,  76,  161,  165,  341,  348;  pis. 

20,  24. 
acervata,  Asterias  13,  36,  59,  107,  108, 

338;  pis.  27,  106. 
acervata  borealis,  Asterias  370. 
acutispina,  Coscinasterias  38,  46,  373. 
Adelasterias,  new  genus  352,  360,  371, 

374- 

Adelasterias  papillosa  360,  372. 
Adetopneusia  24. 
cequalis,  Asterias  128. 
aequalis,  Leptasterias  36,  128;  pis.  25, 

56. 
aequalis,  Mediaster,  291,  296,  343,  344, 

346,  348 ;  pis.  2,  3,  5. 
aequalis,  var.  compacta,  Leptasterias, 

new  variety  61,  130,  341,  344;  pi. 

16. 
aequalis,  var.  concinna,  Leptasterias, 

new  variety  61,  132,  344. 
aequalis,  var.  nana,  Leptasterias,  new 

variety  61,  132,  341,  344,  345. 
affinis,  Asterias  259,  261. 
affinis,  Crossaster  261. 
affinis,  Pteraster  278. 
alaskensis,  Leptasterias  epichlora,  new 

subspecies  13,  39, 61,  in,  136,  338, 

341;  pis.  16,  28,  84. 
alba,  Cosmasterias  358. 
albertensis,  Parasterias,  new  species 

S3,  55,  187.  342,  348;  pis.  57,  70. 
alboverrucosa,  Asterias  259,  261,  262. 
albula,  Asterias  41. 


albula,  Stephanasterias  39,  40,  41,  55, 

67,  147,  338. 

albulus,  Asteracanthion  147. 
albulus,  Nanaster  41. 
albulus,  Stichaster  41,  147. 
aleutica,  Henricia  213. 
Alexandraster  205. 
Alexandraster  inflatus  206. 
alexandri,  Coscinasterias  350,  373. 
alexandri,  Stolasterias  372. 
Allasterias  38,  53,  55,  66,  188,  352. 
Allasterias  amurensis  373. 
Allasterias  anomala  67,  193,  338,  347 ; 

pis.  59,  60,  69,  78. 
Allasterias    forficulosa,    new    species 

67,  194.  373J  Pis.  77,  83,  84. 
Allasterias  migrata  373. 
Allasterias  rathbuni  53,  55, 66, 67, 189, 

337,  338,  347- 
Allasterias  rathbuni  nortonensis  67, 

191,  338,  3475  pL  78. 
Allasterias  versicolor  196,  373. 
Allopatiria  ocellifera  263. 
Allostichaster,  new  genus  362,  363, 

373,  374- 

Allostichaster  polyplax  373. 
alveolata,  Evasterias  troschelii,   new 

variety  63,  157,  162,  341 ;  pi.  62. 
Amphiaster  294. 

Amphiaster  insignis  294,  373;  pi.  98. 
amurensis,  Allasterias  373. 
amurensis,  Asterias  194. 
Anasterias  8,  352,  353,  354. 
Anasterias  belgicae  51,  355,  371. 
Anasterias  chirophora  51,  355. 
Anasterias  lactea  355. 
Anasterias  lysasteria,  new  name  354, 

355,  37i. 

Anasterias  minuta  354,  355,  371. 
Anasterias  perrieri  51,  354,  355. 
Anasterias  studeri  355,  371. 
Anasterias  tenera  355,  371. 
Anasterias  verrillii  354,  355,  371. 

(389) 


390 


INDEX 


angulosa,  Asterias  370. 
annulata,  Labidastrella  373. 
annulata,  Labidiaster  352,  373,  374. 
anomala,  Allasterias  67,  193,  338,  347 ; 

pis.  59,  60,  69,  78. 
anomalus,  Glyphaster  316,  328,  340, 

347 ;  pis.  i,  6. 

Antarctic  species  2,  51,  350,  351. 
antarctica,  Porania  368,  372. 
atitarctica,    Sporasterias   51,   53,   353, 

354,  371. 

antarcticus,  Leptychaster  369. 
antarcticus,  Pedicellaster  367,  371. 
Anthenea  286,  287. 
Anthenea  granulif  era  288. 
Antheneidce,  282,  299. 
Antheneinae  283. 
antillarum,  Henricia  210,  370. 
antonioensis,  Echinaster  363,  372. 
aporus,  Pteraster  277,  278. 
aporus,  Pterasterides  278,  339,  347. 
Ar chaster  arcticus  326. 
Archaster  bairdii  298. 
Archaster  dawsoni  312. 
Archasteridae  283. 
arctica,  Asterias  120. 
arctica,   Henricia,   new   species   239, 

339,  347;  pl.  87. 
arctica,  Leptasterias  120,  338,  347;  pis. 

56,  71,  72,  83. 
arctica,  Tosia  292. 
arcticus,  Archaster  326. 
arcticus,  Astropecten  326. 
arcticus,  Ceramaster  292. 
arcticus,  Leptoptychaster  326. 
arcticus,  Leptychaster  326,  339. 
arcticus,  Solaster  dawsoni  347. 
arcticus,  Tosiaster  292,  339;  pis.  50, 

99. 

arcuatus,  Pteraster  tesselatus  269. 
armatus,  Astropecten  373. 
armatus,  Astropecten  317,  319. 
aspera,  Henricia  241,  339,  342,  346, 

348. 

Asteracanthion  101,  114. 
Asteracanthion  albulus  147. 
Asteracanthion  clavatum  358. 
Asteracanthion  fulvum  358. 
Asteracanthion  katherina  112. 
Asteracanthion  margaritifer  69. 


Asteracanthion  mite  358. 
Asteracanthion  problema  147  • 
Asteracanthion  rubens  373. 
Asteracanthion  spectabile  358. 
Asteracanthium  camtschaticum  114. 
Asterias  4,  n,  16,  17,  18,  33,  34,  43, 

44.  55,  58,  68,  ioi,  352. 
Asterias  acervata  13,  36,  59,  107,  108, 

338;  pis.  27,  106. 
Asterias  acervata  borealis,  new  name 

107,  in,  370. 
Asterias  aqualis  128. 
Asterias  afKnis  259,  261. 
Asterias  alboverrucosa  259,  261,  262. 
Asterias  albula  41. 
Asterias  amurensis  194. 
Asterias  angulosa  370. 
Asterias  arctica  120. 
Asterias  aurantiaca,  362,  372. 
Asterias  borealis  107,  113. 
Asterias  brachiata  152,  156. 
Asterias  brandti  371. 
Asterias  brevispina  77. 
Asterias  camtschatica   no,   114,   116, 

127,  132,  135. 
Asterias  capitata  81. 
Asterias  conferta  73,  77- 
Asterias  (Cosmasterias)  tomidata  48. 
Asterias  cribraria  148. 
Asterias  crispatus  330. 
Asterias  cunninghami  353. 
Asterias  decemradiatus  254. 
Asterias  (Diplasterias)  epichlora  152. 
Asterias  douglasi  107,  113,  114. 
Asterias  dubia  97. 
Asterias  echinophora  207. 
Asterias  endeca  244. 
Asterias  epichlora  132,  158. 
Asterias  exquiseta  83. 
Asterias  fissispina  76. 
Asterias  forbesi  3,  5,  10,  18,  38,  188, 

370. 

Asterias  forreri  47,  179. 
Asterias  gelatinosa  357. 
Asterias  gigantea  89,  97. 
Asterias  granularis  290. 
Asterias  helianthoides  198. 
Asterias  hexactis  126. 
Asterias  hyadesi  354. 
Asterias  janthina  69,  73. 


INDEX 


391 


Asterias  katherinae   13,  36,   59,    112, 

340,  344,  348;  pis.  51,  83. 
Asterias  lurida  337,  346. 
Asterias  lutkenii,  44,  82,  83. 
Asterias  meridionalis  361,  372. 
Asterias  militaris  272. 
Asterias  miniata  264. 
Asterias  minuta-  107. 
Asterias  multiclava  59,  no,  114,  338, 

347;  pis.  58,  59,69,  84. 
Asterias  nanimensis,  new  species  59, 

105,  340;  pl-  61. 
Asterias  neglecta  358. 
Asterias  obtusispinosa  358. 
Asterias  ochracea  69,  77,  82. 
Asterias  oculata  226. 
Asterias  papillosa  372. 
Asterias  papposa  259. 
Asterias  paucispina  98. 
Asterias  perrieri  354,  356. 
Asterias  pertusa  226. 
Asterias  planci  364. 
Asterias  Polaris  107,  108,  113,  370. 
Asterias  polythela  59,  i<>4»  338,  347; 

pis.  55,  70,  72,  79,  84. 
Asterias  rosea  40,  362. 
Asterias  rubens  44,  55,  68,  188,  337, 

346. 

Asterias  rubens  107,  189. 
Asterias  rugispina  51,  53,  353,  355- 
Asterias  rupicola  356. 
Asterias  rustica  357. 
Asterias  saanichensis  132,  135. 
Asterias  sanguinolenta  226. 
Asterias  sertulifera  100. 
Asterias  spirabilis  51,  354. 
Asterias  spitsbergensis  148,  150. 
Asterias  steineni  361. 
Asterias  stellionura  51. 
Asterias  (Stichaster)  polyplax  125. 
Asterias  studeri  51. 
Asterias  sulcifera  48,  358,  361. 
Asterias  troschelii  114,  151,  156. 
Asterias  turqueti  372. 
Asterias  (Urasterias*)  forcipulata  180. 
Asterias  vancouveri  13,  125. 
Asterias  varia  51. 
Asterias  victoriana  59,  102,  340,  348; 

pis.  53,  54,  69,  82. 
Asterias  violacea  107. 


Asterias  vulgaris  38,  370. 
Asteriidae  27,  30,  39,  42,  43,  56. 
Asteriida,  Sladen's  43. 
Asteriinae  26,  39,  42,  43,  49,  50. 
Asterina  262. 
asterina,  Cycethra  366. 
Asterina  gibbosa  262,  263,  371. 
Asterina  miniata  264. 
Asterina  minuta  262. 
Asterina  perrieri  365,  372. 
Asterina  pygmcea  371. 
Asterinidae  204,  262. 
Asterinides  folium  263,  371. 
Asterinides  modesta  373. 
Asterinides  perrieri  365,  372. 
asterinoides,  Calvasterias  357,  371. 
Asterinopsis  penicillaris  263,  373. 
Asterioidea,  classification  24. 
Asterioidea,  derivation  of  name  20. 
Asteriscus  263. 
asteriscus,  Cycethra  366. 
Asteriscus  miniatus  264. 
Asterodon  belli  367. 
Aster odon  granule sus  367. 
Asterodon  singularis  367,  372. 
Asteropidse  282,  284,  304. 
Asteropinae,  new  subfamily  305. 
Asteropsis  305. 
Asteropsis  imbricata  306. 
asthenactis,  Henricia  213. 
asthenosoma,  Luidia  336,  345,  346. 
Astrogonium  287,  289,  300. 
Astrogonium  granulare  290. 
Astrogonium  granularis  287. 
Astrogonium  patagonicus  368. 
Astropecten  281,  316,  317,  348. 
Astropecten  arcticus  326. 
Astropecten  armatus  373. 
Astropecten  armatus  317,  319. 
Astropecten    californicus    319,    320, 

345,  346;  pis.  50,  TOO,  101,  102. 
Astropecten  ctenophorus  321. 
Astropecten  erinaceus  347. 
Astropecten  -derstedii  318,  319,  346,  373. 
Astropecten    ornatissimus    320,    346, 

349- 
Astropecten    siderealis,    new    species 

317.  345,  346,  349;  pl-  50. 
Astropectinidae  283,  285,  314. 


392 


INDEX 


Astropectinides,  new  genus  316,  317, 

331,  372,  374. 
Astropectinides   mesacutus   316,   321, 

372. 

atlantica,  Tonia  40,  362. 
attenuate,    Henricia    leviuscula    217, 

218,  342,  344. 

aurantiaca,  Asterias  362,  372. 
aurantiacus,  Stichaster  40,  362. 
australis,  Coelasterias  373. 
australis,  Ctenodiscus  333,  369. 
australis,  Pisaster  liitkeni,  new  variety 

85,  88,  345. 

australis,  Solaster  366,  368,  372. 
austro-granularis,   Ceramaster  369. 
austro-granularis,  Pentagonaster  369. 
Avelata,  suborder  204. 

bairdii,  Archaster  298. 

bairdii,  Isaster  298. 

bairdii,  Mediaster  298,  371 ;  pis.  2,  3. 

belgicae,  Anasterias  51,  355,  371. 

belli,  Asterodon  367. 

bellonae,  Luidia  367. 

bellula,  Callopatiria  263. 

Benthopectinidae  283,  310. 

Benthopectinida  311. 

Benthopectininae  283,  310,  311. 

Beringian  fauna  338,  347. 

Bibliography  374-388. 

bifascialis,  Phataria    (Linckia)    uni- 
fascialis  var.  309. 

biordinate,  Qrthasterias,  new  species 
48,  64,  172,  341 ;  pis.  63,  82. 

biserialis,  Peribolaster  366. 

biserialis,  Phataria  346. 

biseriatus,  Granaster  360,  362,  372. 

biseriatus,   Hemiasterias,   new   genus 
362,  372. 

bispinosa,  Patiriella  364,  372. 

Blakiaster  322,  324. 

Blakiaster  conicus  322,  371. 

borealis,  Asterias  107,  113. 

borealis,  Asterias  acervata  107,  in 
370. 

borealis,  Henricia  235,  236,  239. 

borealis,  Henricia  tumida,  new  sub- 
species 336,  339,  342,  347;   pis. 
12,  86,  88. 
brachiata,  Asterias  152,  156. 


Brachiolaria,  larval  form  24. 
bradleyi,  Mithrodia  373 ;  pi.  107. 
brandti,  Asterias  371. 
brandti,  Cosmasterias  360,  372. 
brasiliensis,  Echinaster  363. 
brasiliensis,  Enoplopatiria  263. 
brevispina,  Asterias  77. 
brevispina,  Luidia  347. 
brevispina,  Pisaster  77- 
brevispinus,  Pisaster  57,  77,  93,  340, 

344,  345  J  Pis.  41,  44,  45,  69,  76. 
briareus,  Coronaster  49,  370. 
briareus,  Thy  one  10. 
Brisingidae  15,  24,  26,  374. 
Brisinginae  26. 

Brooding  of  the  young  39. 
Bunodaster  316,  331. 
Bunodaster  ritteri  316,  333,  345,  348; 
pis.  86,  104,  105. 

calcarata,  Patiriella  364,  365,  372. 
Californian  fauna  337,  343. 
californica,  Orthasterias,  new  species 

48,  64,  174,  185,  344,  350;  pis.  68, 

70,  80,  81. 
californicus,   Astropecten   319,   320, 

345,  346;  pis.  50,  ioo,  101,  102. 
californicus,  Rathbunaster  197. 
Callopatiria  bellula  263. 
Callopatiria  obtusa  373. 

calva,  Cycethra  366. 
Calvasterias  42,  353,  357. 
Calvasterias  asterinoides  357,  371. 
Calvasterias   stolidota  42,  354,    357, 

37L 
camtschatica,  Asterias  no,  114,  116, 

127,  132  135- 

camtschaticum,   Asteracanthium    114. 
candicans,  Coscinasterias  367. 
candicans,  Stolasterias  367. 
Cannibalism  among  starfishes  4. 
capensis,  Pteraster  278. 
capitata,  Asterias  81. 
capitatus,  Pisaster  19,  57,  81,  344,  3455 

pis.  36,  56. 
carinella,  Leptasterias  epichlora,  new 

variety  61,  137,  341- 
carribaeus,  Pteraster  278. 
Ceramaster  389. 


INDEX 


393 


Ceramaster  arcticus  292. 
Ceramaster  austro-granularis  369. 
Ceramaster  granularis  290,  293,  339, 

343 ;  pis.  4,  50. 
Ceramaster  japonicus  292. 
Ceramaster  leptoceramus  292. 
Ceramaster  patagonicus  291,  339,  343, 

347,  349,  368. 
Chaetasteridae  283. 
Cheiraster  gerlachei  369. 
Cheirasteridce  311. 
chelifera,  Distolasterias,  new  species 

66,  185,  342;  pis.  81,  no. 
chilensis,  Patiria  348,  364,  372. 
chirophora,  Anasterias  51,  355. 
chirophora,   Paedasterias,  new  genus 

355,  371. 

Chitonasterinas  283. 
Cladaster  300. 
clarki,  Henricia  213. 
clathrata,  Luidia  7,  371 ;  pi.  103. 
clavatum,  Asteracanthion  358. 
coccinea,  Patiria  263,  373. 
cocosana,  Sporasterias  350. 
coei,  Leptasterias,  new  species  36,  61, 

123,  178,  340,  348 ;  pis.  9,  17. 
Coelasterias  40. 
Ccelasterias  australis  373. 
cognatus,  Mimaster  368. 
columbise,  Linckia  309,  337,  349. 
columbiana,    Orthasterias,    new    spe- 
cies n,  48,  50,  64,  1 68,  178,  183, 

341,  344,  346,  348;  pis.  24,  35,  65, 

78,  79,  109. 
compacta,  Leptasterias  sequalis,  var. 

61,  130,  341,  344. 
compta,  Leptasterias  8,  121,  370. 
concinna,   Leptasterias   aequalis,   new 

variety  61,  132,  344. 
conferta,  Asterias  73. 
confertus,  Pisaster  57,  73,  167,  340; 

pl.  38. 

conicus,  Blakiaster  322,  371. 
constellatus,  Solaster  257,  343,  348; 

pis.  46,  90,  91,  93,  94. 
corniculatus,  Ctenodiscus  331. 
Coronaster  30. 
Coronaster  briareus  49,  370. 
Coscinasterias  33,  34,  36,  38,  39,  43, 


Coscinasterias  acutispina  38,  46,  373. 
Coscinasterias  alexandri  350,  373. 
Coscinasterias  candicans  367. 
Coscinasterias  echinata  46. 
Coscinasterias  gemmifera  46. 
Coscinasterias  jehennesii  46. 
Coscinasterias  muricata  46,  49,  373. 
Coscinasterias      (Stolasterias)      ten- 

uispina  46,  49. 
Coscinasterias  tenuispina  38,  43,  45, 

370. 

Cosmasterias  33,  44,  48,  353,  3p8. 
Cosmasterias  67. 
Cosmasterias  alba  358. 
Cosmasterias  brandti  360  372. 
Cosmasterias  fernandensis  360,  372. 
Cosmasterias  lurida  36,  48,  50,  358, 

37i. 

Cosmasterias  polygrammus  360,  371. 
Cosmasterias   sulcifera  48,   358,  360, 

361,  371. 
Cosmasterias   tomidata  48,   50,   359, 

371- 

crassa,  Henricia  leviuscula  217. 
crassa,  Parapatiria  263. 
crassus,    Odontaster    304,    345,   346, 

349- 

cremens,  Odontaster  368. 
Cribella  209. 
Cribella  hyadesi  363. 
Cribella  oculata  226. 
Cribella  rosea  226. 
cribraria,  Asterias  148. 
cribraria,    Ctenasterias    53,   60,    148, 

338;  pl.  25. 

cribraria,  Leptasterias  8. 
Cribraster  sladeni  363,  372. 
Cribrella  211. 
Cribrella  leviuscula  215. 
Cribrella  oculata  226. 
Cribrella  pectinata  230. 
Cribrella  prastans  368. 
Cribrella  sanguinolenta  223,  226. 
Cribrella  simplex  368. 
Cribrella  spiculifera  220. 
Cribrellae  129. 
cribrosus,  Pteraster  278. 
crispatus,  Asterias  330. 
crispatus,  Ctenodiscus  330,  340,  343, 

345,  347,  3495  pis.  5,  49- 


394 


INDEX 


Crossaster  13,  15,  358. 
Crossaster  243. 
Crossaster  affinis  261. 
Crossaster  korenii  261,  371. 
Crossaster   papposus   249,   259,   339, 

343;  pis.  5,8,9,49- 
Cryptasterias,   new  genus   362,   372, 

374- 

Cryptasterias  turqueti  362,  372. 
Cryptozonia  24. 
Ctenaster  263. 
Ctenasterias,  new  genus  53,  55,  148, 

374- 
Ctenasterias    cribraria    53,    60,    148, 

338;  pi.  25. 
Ctenasterias    spitzbergensis,    53,    55, 

148,  270. 

Ctenodiscinae  283,  329. 
Ctenodiscus,  315,  329,  330. 
Ctenodiscus  australis  333,  369. 
Ctenodiscus  corniculatus  331. 
Ctenodiscus   crispatus   330,  340,   343, 

345,  347,  3495  pis.  5,  49- 
Ctenodiscus  krausei  331. 
Ctenodiscus  polaris  330. 
Ctenodiscus  procurator  333. 
Ctenodiscus  pygmceus  330. 
ctenophorus,  Astropecten  321. 
Culcita  288. 

cunninghami,  Asterias  353. 
cunninghami,  Sporasterias  353. 
Cycethra  asterina  366. 
Cycethra  asteriscus  366. 
Cycethra  calva  366. 
Cycethra  electilis  366. 
Cycethra  elongata  366. 
Cycethra  ganeriodes  366. 
Cycethra  lahillei  366,  372. 
Cycethra  media  366. 
Cycethra  nitida  366. 
Cycethra  pinguis  366. 
Cycethra  regularis  366. 
Cycethra  simplex  366,  372. 
Cycethra  subelectilis  366. 
Cycethra  verrucosa  366,  372. 
Cystidea,  number  of  rays  15. 

danae,  Pteraster  278. 
dawsoni,  Acantharchaster  312. 
dawsoni,  Archaster  312. 


dawsoni  arctica,   Solaster,  new   sub- 
species 252,  253,  347;  pi.  87. 
dawsoni,  Luidiaster  312,  339,  343,  349 ; 

pis.  33,  34,  35- 
dawsoni,  Orthasterias,  new  species  48, 

65,  172,  175,  342;  pis.  23,  75,  80, 
81. 

dawsoni,  Solaster  349,  339,  342,  348; 

pis.  46,  90,  91,  92. 
decemradiatus,  Asterias  254. 
decemradiatus,  Solaster  254. 
densa,     Evasterias     troschelii,     new 

variety  63,  161,  341. 
densispina,  Henricia  223,  373. 
densus,  Perknaster  364. 
Dermasterias  68. 
Dermasterias  imbricata  14,  306,  343, 

345,  348 ;  pis.  50,  86,  97. 
Dermasterias  imbricata,  var.  valvuli- 

f  era  305,  308,  343 ;  pi.  29. 
Dermasterias  inermis  306. 
Desmopatiria  263. 

Desmopatiria  flexilis  263,  364,  372. 
Development,  39. 
diomedeas,  Hydrasterias  350. 
Diplasterias  43,  44,  101,  358. 
Diplasterias  loveni  361. 
Diplasterias  lutkeni  361. 
Diplasterias  papillosa  360. 
Diplasterias  spinosa  361,  372. 
Diplasterias  steineni  361. 
Diplasterias  turqueti  362. 
diplax,  Linckia  309. 
Diplopteraster  203,  267,  273,  279. 
Diplopteraster  multipes  279,  339. 
Diplopteraster  peregrinator  368. 
Diplopteraster  verrucosus  368. 
dispar,  Leptasterias,  new  species  63, 

115,  142,  341;  pi.  16. 
Distolasterias  33,  47,  66,  185,  350. 
Distolasterias  chelifera,  new  species, 

66,  185,  342;  pis.  81,  no. 
Distolasterias  stichantha  44,  49,   185, 

373- 

Distribution,  geographical  18,  337. 
Dorsal  ossicles,  form  34. 
Dorsal  spines,  form  34. 
douglasi,  Asterias  107,  113  114. 
dubia,  Asterias  97. 


INDEX 


395 


dyscrita,  Henricia  leviuscula  223,  342, 

344- 
Dytaster  285. 

Echinaster  101,  206. 

Echinaster  209. 

Echinaster  antonioensis  363,  372. 

Echinaster  brasiliensis  363. 

Echinaster  eschrichtii  226. 

Echinaster  lepidus  363. 

Echinaster  oculatus  226. 

Echinaster    (Othilia)    robustus,   new 

species  207. 

Echinaster  (Othilia)  tenuispinus  346. 
echinaster,  Poraniopsis  349,  363,  372. 
Echinaster  (Rhopia)  lepidus  372. 
Echinaster  robustus  342. 
Echinaster  sanguinolenta  226. 
Echinaster  sarsii  226. 
Echinaster  scrobiculata  227. 
Echinaster  sentus  207. 
Echinaster  smithii  368. 
Echinaster  Solaris  364. 
Echinaster  spinosus  207. 
Echinaster  tenuispinus  208,  346,  349, 

373  J  pl-  107. 

Echinasteridae  204,  205. 
Echinasterine  205,  242. 
echinata,  Coscinasterias  46. 
echinites,  Acanthaster  364,  373. 
echinophora,  Asterias  207. 
echinus,  Acanthaster  364. 
Eggs  and  young  7,  9,  n. 
electilis,  Cycethra  366. 
ellisii,  Acanthaster  364,  373. 
elongata,  Cycethra  366. 
endeca,  Asterias  244. 
endeca,  Solaster  n,  243,  244,  248,  249, 

339;  pis.  9,  87,  89. 
Enoplopatiria  brasiliensis  263. 
Enoplopatiria  marginata  263,  365,  371. 
Enoplopatiria    siderea   365,    373;    pl. 

109. 

epichlora,  Asterias  132,  156. 
epichlora,  Asterias  (Diplasterias)  152. 
epichlora  alaskensis,  Leptasterias,  new 

subspecies  13,  39,  61,  in,  136,  338, 

341 ;  pis.  16,  28,  84. 


epichlora    alaskensis,    var.    carinella, 

Leptasterias,  new  variety  61,  137, 

341 ;  pl.  16. 
epichlora  alaskensis,  var.  siderea, 

Leptasterias,  new  variety  61,  137, 

341 ;  pl.  16. 
epichlora,  Leptasterias  8,  17,  36,  60, 

62,  no,  132,  341,  348. 
epichlora  miliaris,  Leptasterias,  new 

subspecies  60,  61,  138,  341. 

epichlora  plena,  Leptasterias,  new 
subspecies  61,  140,  341. 

epichlora  pugetana,  Leptasterias,  new 
subspecies  61,  142,  341. 

epichlora,  var.  regularis,  Leptasterias, 
new  variety  63,  139,  341. 

epichlora,  var.  subnodulosa,  Leptas- 
terias, new  variety  62,  73,  139, 

341- 

epichlora,  var.  subregularis,  Leptas- 
terias, new  variety  63,  139,  341. 

equestris,  Goniaster  286. 

equestris,  Hippasteria  286. 

erinaceus,  Astropecten  347. 

eschrichtii,  Echinaster  226. 

eschrichtii,  Henricia  212,  236,  329. 

eustyla,  Orthasterias  168. 

Evasterias,  new  genus  28,  33,  51,  54, 

63,  151,  352,  374- 

Evasterias  acanthostoma,  new  species 

11,  64,  76,  161,  165,  341,  348;  pis. 

20,  24. 
Evasterias  troschelii  n,  36,  51,  54,  63, 

151,  34i,  344,  348;  pis.  22,  25,  26, 

106. 
Evasterias   troschelii,   var.   alveolata, 

new  variety  63,  157,  162,  341 ;  pl. 

62. 
Evasterias  troschelii,  var.  densa,  new 

variety  63,  161,  341. 
Evasterias  troschelii,  var.  parvispina, 

new  variety  63,  163,  341. 
Evasterias  troschelii,  var.  rudis,  new 

variety  63,  158,  341. 
Evasterias  troschelii,  var.  subnodosa, 

new  variety  63,  163,  341. 
exquiseta,  Pisaster  liitkenii  345. 
exquiseta,  Asterias  83. 
Eye  or  eye-spot  of  starfishes  9. 


39^ 


INDEX 


falklandica,  Ganeria  365,  372. 

Faunal  relations  and  distribution  18. 

f  elipes,  Stichaster  49. 

fernandensis,  Cosmasterias  360,  372. 

fernandensis,  Polyasterias  360,  371. 

fimbriata,  Patiriella  364  365,  372. 

fissispina,  Asterias  76. 

fissispina,  Pisaster  76. 

fissispinus,  Pisaster  34,  57,  76,  344;  pi. 

39- 

flavescens,  Hosea  287. 
Aeuriaisi,  Goniopecten  369. 
fleuriaisi,  Psilaster  369. 
flexilis,  Desmopatiria  263,  364,  372. 
foliata,  Luidia  11,  334,  343,  345,  346; 

pis.  100,  103,  105. 
foliculata,   Peribolaster  366,  372. 
folium,  Asterinides  263,  371. 
Food  and  feeding  habits  3,  16. 
forbesi,  Asterias  3,  5,  10,  17,  38,  188, 

370. 

Forcipulata  24,  41. 
forcipulata,     Asterias     (Urasterias) 

180. 

forcipulata,  Orthasterias  n. 
forcipulata,   Orthasterias   forreri  65, 

1 80,  342 ;  pis.  62,  70,  88. 
Forcipulosa,  order  24,  26. 
forficulosa,  Allasterias,   new   species 

67,  194,  3735  pis.  77,  83,  84. 
forreri,  Asterias  47, 179. 
forreri   forcipulata,   Orthasterias   65, 

1 80,  342;  pi.  62,  70,  88. 
forreri,  Orthasterias  19,  65,  179,  344, 

345;  pis.  65,  66,  70,  77,  80. 
forreri,    Orthasterias     (Stylasterias) 

342. 

forreri,  Stylasterias  48,  50. 
Fossil  starfishes  2,  12,  13,  14,  20,  197. 
franciscus,  Linckia  211. 
fulvum,  Asteracanthion  358. 
fuscus,  Perknaster  364. 

galapagensis,  Sporasterias  350. 
galaxides,  Solaster  248,  342,  348;  pis. 

46,  87,  89. 
Ganeria  falklandica  365,  372. 


Ganeria  hahni  365. 

Ganeria  papillosa  365,  372. 

Ganeria  robusta  365. 

Ganeriidae  204. 

ganeriodes,  Cycethra  366. 

Gastraster  studeri  360,  372. 

gayi,  Patiria  364,  372. 

gelatinosa,  Asterias  357. 

gelatinosus,  Meyenaster  54,  357,  371. 

gemmifera,  Coscinasterias  46. 

Genital  pores  39. 

Geographical  distribution  18,  337-352. 

gerlachei,  Cheiraster  360. 

gibber,  Retaster  366,  372. 

gibbosa,  Asterina  262,  263,  371. 

gigantea,  Asterias  89,  97. 

giganteus,  Pisaster  n,  57,  84,  87,  89, 
98,  344;  pi.  37. 

glaber,  Porania  368. 

glacialis,  Marthasterias  46,  47,  49,  370. 

Glyphaster  316,  325,  337. 

Glyphaster  anomalus  316,  328,  340, 
347 ;  pis.  i,  6. 

Gnathaster  303. 

Gnathaster  grayi  367. 

Gnathaster  pilulatus  367. 

Gnathasterituz  283,  302. 
Goniaster  300. 
Goniaster  equestris  286. 
Goniaster  granularis  290. 
Goniaster  nodosus  286. 
Goniaster  obtusangulus  287. 
Goniaster  reticulatus  286. 
Goniaster  tesselatus  286. 
Goniasteridae  281,  282,  285,  286. 
Goniasterinae  283,  289. 
Goniodiscides  sebce  288. 
Goniodiscus  288. 
Goniodiscus  pentagonulus  287. 
Goniopecten  fleuriaisi  369. 
Goniopectinidae  283. 
gonolena,    Orthasterias,   new   species 
65,  184,  344,  345,  3495  pis.  67,  68, 
69,  82. 

gracilis,  Pteraster  271,  343,  348. 
gracilis,  Stephanasterias  146. 
Granaster  40,  353,  362. 
Granaster  biseriatus  360,  362,  372. 
Granaster  nutrix  372. 
grandis,  Pisaster  13. 


INDEX 


397 


granulare,  Astrogonium  290. 
granularis,  Asterias  290. 
granularis,  Astrogonium  287. 
granularis,  Ceramaster  290,  293,  339, 

343 ;  pis.  4,  50. 
granularis,  Goniaster  290. 
granularis,  Pentagonaster  290. 
granularis,  Tosia   (Ceramaster)   290. 
granulif  era,  Anthenea  288. 
granulosus,  Asterodon  367. 
granulosus,  Odontaster  367,  372. 
grayi,  Gnathaster  367. 
grayi,  Odontaster  367,  372. 
grayi,  Pisaster  58,  91,  97,  344. 
guernei,  Sclerasterias  44,  50,  370. 
guildingii,  Linckia  309,  310,  337,  346. 
gunneri,  Urasterias  52. 
Gymnasteriidce  304. 

hahni,  Ganeria  365. 
hebes,  Pteraster  270,  343 ;  pi.  96. 
helianthoides,  Asterias  198. 
helianthoides,  Pycnopodia  II,  35,  198, 

342,  344,  348;  pis.  29,  30,  31,  73, 

74,88. 

helianthus,  Heliaster  372. 
Heliaster  12,  13,  15,  40. 
Heliaster  helianthus  372. 
Heliasterinae  26,  39,  40. 
Hemiasterias,  new  genus  362,  374. 
Hemiasterias  biseriatus  362,  372. 
Henricia  8,  17,  18,  144,  209. 
Henricia  aleutica  213. 
Henricia  antillarum  210,  370. 
Henricia  arctica,  new  species  239,  339, 

347;  Pi-  87. 
Henricia  aspera  241,  339,  342,  346, 

348. 

Henricia  asthenactis  213. 
Henricia  borealis  235,  236,  239. 
Henricia  clarki  213. 
Henricia  densispina  223,  373. 
Henricia  eschrichtii  212,  236,  239. 
Henricia  heteractis  210,  373. 
Henricia  hyadesi  363,  368,  372. 
Henricia  japonica  223,  373. 
Henricia  leviuscula  14,  213,  215,  338, 

342,  344,  346 ;  pis.  12,  13. 
Henricia   leviuscula,    var.    annectens 

224,  344,  346. 


Henricia    leviuscula,    var.    attenuata 

217,  218,  342,  344. 

Henricia  leviuscula,  var.  crassa  217. 
Henricia     leviuscula,     var.     dyscrita 

223,  342,  344. 

Henricia  leviuscula,  var.  inequalis, 
new  variety  219,  342;  pi.  88. 

Henricia  leviuscula,  var.  leviuscula 
217,  342. 

Henricia  leviuscula,  var.  lunula,  new 
variety  213,  218,  342,  344;  pi.  88. 

Henricia   leviuscula,   var.   multispina 

220,   222,   338,   342. 

Henricia     leviuscula,    var.   pectinata 

229,  230. 
Henricia  leviuscula,  var.  spatulifera, 

224,  342,344,  346;  pl.  5,  14- 
Henricia  leviuscula  spiculifera   220, 

338,  342 ;  pis.  87, 107. 

Henricia  longispina  342,  348. 

Henricia  obesa  363,  372. 

Henricia  oculata  226. 

Henricia  pagenstecheri  363,  372. 

Henricia  pectinata  212. 

Henricia  polyacantha  213. 

Henricia  praestans  368. 

Henricia   sanguinolenta    n,    14,   212, 

226,  338,  342 ;  pis.  49,  88. 
Henricia  sanguinolenta  miliaris,  new 

subspecies  234,  370;  pl.  88. 
Henricia  sanguinolenta,  var.  pectinata 

229,  230,  338;  pl.  49. 
Henricia    sanguinolenta,    var.    rudis, 

new  variety  233,  338,  347 ;  pl.  86. 
Henricia  sexradiata  371. 
Henricia  simplex  368. 
Henricia  spatulifera  224. 
Henricia  spiculifera  220. 
Henricia  studeri  363. 
Henricia  tumida  214,  234,  239,  240, 

338,  347  5  pis.  12,  87. 
Henricia  tumida  borealis,  new  sub- 
species 236,  339,  342,  347 ;  pis.  12, 

86,88. 

Henricides,  new  genus  210,  373,  374. 
Henricides  heteractis  210,  373. 
heteractis,  Henricia  210,  373. 
Heterasterias,  new  genus  46,  373,  374. 
Heterasterias  volsellata  27,  47,  49,  373. 


INDEX 


he  fact  is,  Asterias  126, 

hexactis,  Leptasterias  13,  61,  125,  126, 

340,  344,348;  pl-25- 
hexactis,  Pteraster  274. 
hexactis,  Pteraster  (Temnaster)  274. 
hexactis,  Temnaster  267,  274. 
Hexaster  267. 

Hexaster  obscurus  274,  275. 
Hippasteria  300. 
Hippasteria  equestris  286. 
Hippasteria  hyadesi  368. 
Hippasteria  phrygiana  286,  301,  371 ; 

pis.  47,  48,  49. 
Hippasteria  spinosa  301,  339,  343,  345, 

346,  348;  Pis.  SO,  98. 
Hippasterias  Bell  300. 
Hippasteriinae  283,  299. 
hispidus,  Odontaster  349. 
Hosea  287. 
Hosea  flavescens  287. 
hyadesi,  Asterias  354. 
hyadesi,  Cribella  363. 
hyadesi,  Henricia  363,  368,  372. 
hyadesi,  Hippasteria  368. 
hyadesi,  Sporasterias  354. 
Hybrids  or  freaks  17,  18. 
Hydrasterias  43,  44,  350. 
Hydrasterias  diotnedeae  350. 
Hymenaster  267^ 
Hymenaster  obscurus  267. 
Hymenaster  perspicuus  368. 
Hymenasterinae,   new  subfamily  266, 

267. 

Ilyaster  315. 

imbricata,  Aster  op  sis  306. 

imbricata,  Dermasterias  14,  306,  343, 
345,  348;  pis.  29,  50,  86,  97. 

imbricata,  var.  valvulifera,  Dermas- 
terias 305,  308,  343. 

Incubation  7. 

inequalis,  Henricia  leviuscula,  new 
variety  219,  342;  pi.  88. 

inequalis,  Leptasterias,  new  species 
60,  117,  143,  340;  pl.  73- 

inermis,  Dermasterias  306. 

inermis,  Parastropecten  325. 

inflata  flexibilis,  Poraniopsis  206. 

inflata,  Poraniopsis  206,  344,  346,  349, 
363. 


inflatus,  Alexandraster  206. 

ingouffi,  Pteraster  368. 

insignis,  Amphiaster  294,  373;  pi.  98. 

Isaster  295. 

Isaster  bairdii  298. 

janthina,  Asterias  69,  73. 
Japanese  current  2. 
japonica,  Henricia  223,  373. 
japonicus,  Ceramaster  292. 
Jaws  and  oral  spines  36. 
jehennesii,  Coscinasterias  46. 

katherina,  Asteracanthion  112. 
katherinae,  Asterias   13,  36,  59,   112, 

340,  344,  348;  pis.  51,  52,  83. 
katherince,  Asterias  Per.  58,  89,  91,  97. 
kerguelenensis,  Leptychaster  316,  325, 

35i,  369,  372. 
koehleri,    Orthasterias   64,    175,    183, 

342;  pi.  75. 

korenii,  Crossaster  261,  371. 
Korethrasterinae  242. 
krausei,  Ctenodiscus  331. 

Labidastrella,  new  genus  352,  373,  374. 

Labidastrella  annulata  373. 

Labidiaster  12,  15,  26. 

Labidiaster  annulata  352,  373,  374. 

Labidiaster  radiosus  352,  371. 

Labidiasterinae  26. 

lactea,  Anasterias  355. 

lahillei,  Cycethra  366,  372. 

Lahillia  205. 

Lahillia  mira  363. 

Larval  stages  7. 

Lebrunaster  paxillosus  365,  372. 

lebruni,  Pteraster  368. 

Leiaster  308. 

lepidus,  Echinaster  363. 

lepidus,  Echinaster  (Rhopia)  372. 

leptalea,  Leptasterias,  new  species  60, 

119,  340,  348. 
Leptasterias  8,  38,  41,  43,  50,  53,  55, 

56,  60,  72,  116,  351. 
Leptasterias  aequalis  36,  128;  pis.  25, 

56. 
Leptasterias  aequalis,   var.   compacta, 

new  variety  61,  130,  341,  344;  pi. 

16. 


INDEX 


399 


Leptasterias    asqualis,    var.    concinna, 

new  variety  61,  132,  344. 
Leptasterias  asqualis,  var.  nana,  new 

variety  61,  132,  34*,  344,  345- 
Leptasterias  arctica  120,  338,  347 ;  pis. 

56,  71,  72,  83. 
Leptasterias  coei,  new  species  36,  61, 

123,  178,  340,  348;  pis.  9,  17. 
Leptasterias  compta  8,  121,  370. 
Leptasterias  cribraria  8. 
Leptasterias   dispar,  new  species  63, 

115,  143,  34i;  pl-  16. 
Leptasterias  epichlora  8,  17,  36,  60,  62, 

1 10,  132,  341,  348. 
Leptasterias  epichlora  alaskensis,  new 

subspecies   13,  39,  61,   in,   136, 

338,  341 ;  pis.  28,  85. 
Leptasterias  epichlora  alaskensis,  var. 

carinella,   new   variety  61,    137, 

341;  pi.  16. 
Leptasterias  epichlora  alaskensis,  var. 

siderea,  new  variety  61,  137,  341 ; 

pi.  16. 
Leptasterias   epichlora   miliaris,   new 

subspecies  60,  61,  138,  341. 
Leptasterias  epichlora  plena,  new  vari- 
ety, 61,  140,  341 ;  pi.  58. 
Leptasterias  epichlora  pugetana,  new 

subspecies  61,  142,  341. 
Leptasterias  epichlora,  var.  regularis, 

new  variety  63,  139.  34L 
Leptasterias  epichlora,  var  subnodu- 

losa,  new  variety  62,  73,  139,  341. 
Leptasterias  epichlora,  var.  subregu- 

laris,  new  variety  63,  139,  341. 
Leptasterias  hexactis  13,  61,  125,  126, 

340,  344,  348;  pi.  25. 
Leptasterias  inequalis,  new  species,  36, 

60,  117,  143,  340 ;  pl.  73- 
Leptasterias  leptalea,  new  species  60, 

119,340,348;  pi.  18. 
Leptasterias  littoralis  8,  370. 
Leptasterias  macouni,  new  species  13, 

61,  124. 

Leptasterias  macropora  145. 
Leptasterias  mulleri  55,  124. 
Leptasterias  obtecta,  new  species  60, 

144,  338. 

Leptasterias  tenera  8,  370. 
Leptasterias  vancouveri  61,  125,  340. 


leptoceramus,  Ceramaster  292. 

Leptogonasterinae  283. 

leptolena,   Orthasterias,   new   species 

66,  182,  342 ;  pis.  64,  77. 
Leptoptychaster  325,  326. 
Leptoptychaster  arcticus  326. 
Leptostroteria  24. 
Leptychaster  8,  316,  322,  325. 
Leptychaster  antarcticus  369. 
Leptychaster  arcticus  326,  339. 
Leptychaster  kerguelenensis  316,  325, 

351,  369,  372. 

Leptychaster  millespina  326. 
Leptychaster  pacificus  326,  340,  347; 

pi.  74. 

leviuscula,  Cribrella  215. 
leviuscula,  Henricia  14,  213,  215,  338, 

342,  344,  346 ;  pis.  12,  13. 
leviuscula,  Linckia  215. 
leviuscula,   var.   annectens,   Henricia 

224,  344,  346. 
leviuscula,    var.    attenuata,    Henricia 

217,  218,  342,  344. 

leviuscula,  var.  crassa,  Henricia  217. 
leviuscula,    var.     dyscrita,     Henricia 

223,  342,  344;  pi.  12. 
leviuscula,    var.    inequalis,    Henricia, 

new  variety  219,  342;  pi.  88. 
leviuscula,   var.   leviuscula,   Henricia 

217,  342. 
leviuscula,  var.  lunula,  Henricia,  new 

variety  213,  218,  342,  344;  pi.  88. 
leviuscula,    var.    pectinata,    Henricia 

229,  230. 
leviuscula,  var.  spatulifera,  Henricia 

224,  342,  344,  346;  pis.  5,  14. 
leviuscula  spiculifera,  Henricia  220, 

338,  342 ;  pis.  87,  107. 
Linckia  211,  309. 
Linckia  209. 

Linckia  columbiae  309,  337,  349. 
Linckia  dispar  309. 
Linckia  franciscus  211. 
Linckia  guildingii  309,  310,  337,  346 
Linckia  leviuscula  215. 
Linckia  oculata  226. 
Linckia  ornithopus  310. 
Linckia  pertusa  226. 
Linckia  typus  211. 


400 


INDEX 


(Linckia)  unifascialis,  var.  bifascialis, 

Phataria  309. 
Linckia  variolata  211. 
Linckiadce  282,  308. 
linckii,  Urasterias  33,  51,  55,  66,  181, 

187,  370;  pi.  70. 
Linckiidae  41,  308. 
littoralis,  Leptasterias  8,  370. 
longispina,  Henricia  342,  348. 
Lophaster  pentactis  368. 
Lophaster  stellans  366,  368,  372. 
loveni,  Diplasterias  361. 
loveni,  Podasterias  361. 
ludwigi,  Luidia  336,  345,  346. 
liitkeni,  Diplasterias  361. 
liitkeni,  Podasterias  44,  51,  351,  361, 

372. 

liitkenii,  Asterias,  44,  82,  83. 
liitkenii,  Pisaster  n,  57,  68,  83,  344, 

345,  362 ;  pi.  40. 
liitkenii,  var.  australis,  Pisaster,  new 

variety  85,  88,  345. 

lutkenii,  var.  exquiseta,  Pisaster  345. 
Luidia  6,  281,  315,  334,  348. 
Luidia  asthenosoma  336,  345,  346. 
Luidia  bellonae  367. 
Luidia  brevispina  347. 
Luidia  clathrata  7,  371 ;  pi.  103. 
Luidia  foliolata  n,  334.  343,  345,  346; 

pis.  100,  103,  105. 
Luidia  ludwigi  336,  345,  346. 
Luidia  magellanica  367,  372. 
Luidia  phragma  367. 
Luidiaster  311. 
Luidiaster  dawsoni  312,  339,  343,  349; 

pis.  33,  34,  35- 
Luidiidae  283,  333. 
Luidiina  333. 

lunula,  Henricia  leviuscula,  new  vari- 
ety 213,  218,  342,  344;  pi.  88. 
lurida,  Asterias  337,  346. 
lurida,  Cosmasterias  36,  48,  50,  358, 

371- 
lysasteria,  Anasterias,  new  name  354, 

355,  371. 
Lysasterias  354. 

macouni,  Leptasterias,  new  species  13, 
61,  124. 

macropora,  Leptasterias  145. 


macropora,  Stenasterias  13,  66,  145, 

341,  348;  pis.  50,  74,  84. 
Madreporite  plate  36. 
magelhanica,  Porania  368. 
magellanica,  Luidia  367,  372. 
margaritifera,  Asteracanthion  69.  . 
marginata,  Enoplopatiria  263,  365,  371. 
mariana,  Sporasterias  350. 
marsippus,  Pter aster  273,  339,  347. 
Marthasterias  46,  47,  100,  358. 
Marthasterias  glacialis  46,  47,  49,  370. 
Marthasterias  sertulifera  19,  58,  100, 

345,  349- 

Maternal  instincts  10. 
media,  Cycethra  366. 
Mediaster  285,  295. 
Mediaster  sequalis  291,  296,  343,  344, 

346,348;  pis.  2,  3,  5. 
Mediaster  bairdii  298,  371 ;  pis.  2,  3. 
Mediaster  stellatus  298. 
Mediasterinae  283,  294. 
meridionalis,  Asterias  361,  372. 
meridionalis,  Podasterias  361,  372. 
merriami,   Orthasterias,   new   species 

13,  65,  177,  342;  pis.  18,  19,  75. 
mesacutus,   Astropectinides  316,  321, 

372- 

Meyenaster  54,  353. 
Meyenaster  gelatinosus  54,  357,  371. 
migrata,  Allasterias  373. 
Migratory  habits  2,  4. 
miliaris,  Henricia  sanguinolenta  234, 

370;  pi.  88. 
miliaris,  Leptasterias  epichlora,  new 

subspecies  60,  61,  138,  341. 
militaris,  Asterias  272. 
militaris,  Pteraster  272,  278,  339,  343. 
millespina,  Leptychaster  326. 
Mimaster  cognatus  368. 
Mimasteridse  282. 
miniata,  Asterias  264. 
miniata,  Asterina  264. 
miniata,  Patiria  14,  264,  343,  344,  346, 

347,  348,  349 ;  pi.  7,  108,  109. 
miniatus,  Asteriscus  264. 
minuta,  Anasterias  354,  355,  371. 
minuta,  Asterias  107. 
minuta,  Asterina  262. 
mira,  Lahillia  363. 
mira,  Poraniopsis  363,  372. 


INDEX 


401 


mirabilis,  Tremaster  371. 
mite,  Asteracanthion  358. 
Mithrodia  bradleyi  373 ;  pi.  107. 
Mithrodiidas  204. 
modesta,  Asterinides  373. 
Mollusca,  food  for  starfishes  3. 
Morphological  features  20,  31. 
Mouth  of  starfishes  21. 
mulleri,  Leptasterias  55,  124. 
multiclava,  Asterias  59,  no,  114,  338, 

347 ;  pis.  58,  59,  69,  84. 
multipes,  Diplopteraster  279,  339. 
multipes,  Pteraster  278,  279. 
multipes,  Retaster  279. 
multispina,   Henricia   leviuscula   220, 

338. 

multispinus,  Pteraster  271,  343,  348. 
muricata,   Coscinasterias  46,  49,  373. 
Myonota,  suborder  283,  310. 
Myxasteridae  204. 

nana,  Leptasterias  aequalis,  new  vari- 
ety 61,  132,  34i,  344,  345- 

Nanaster  146. 

Nanaster  albulus  41. 

nanimensis,  Asterias,  new  species  59, 
105,  340;  pi.  61. 

Nectria  287. 

Nectriinae  283. 

neglecta,  Asterias  358. 

neglecta,  Stolasterias  47. 

neglecta,  Stylasterias  48,  370. 

nitida,  Cycethra  366. 

niiida,  Stephanasterias  albula  147. 

nodiferus,  Pisaster  ochraceus  57,  72, 
340,  344;  pi.  56. 

nodosus,  Goniaster  286. 

nodosus,  Oreaster  286. 

nortonensis,  Allasterias  rathbuni  67, 
191,338,347;  pl.  78. 

Notomyota,  order  310. 

nutrix,  Qranaster  372. 

nutrix,  Stichaster  51,  362. 

obesa,  Henricia  363,  372. 
obscurus,  Hexaster  274,  275. 
obscurus,  Hymenaster  267. 
obscurus,  Pteraster  13,  17,  274,  339. 
obscurus,  var.  octaster,  Pteraster  13, 
339,  347- 


obtecta,  Leptasterias,  new  species  60, 

144,  338. 

obtusa,  Callopatiria  373. 
obtusangulus,  Goniaster  287. 
obtusangulus,  Pseudoreaster  287. 
obtusispinosa,  Asterias  358. 
ocellifera,  Allopatiria  263. 
ochracea,  Asterias  69,  77,  82. 
ochraceus,  Pisaster  n,  14,  57,  69,  72, 

340,  344,  345 ;  pl.  21. 
ochraceus,  var.  nodiferus,  Pisaster  57, 

72,  340,  344. 
octaster,  Pteraster  274. 
octoradiatus,  Solaster  368. 
oculata,  Asterias  226. 
oculata,  Cribella  226. 
oculata,  Cribrella  226. 
oculata,  Henricia  226. 
oculata,  Linckia  226. 
oculatus,  Echinaster  226. 
Odontaster  303. 

Odontaster  crassus  304,  345,  346,  349, 
Odontaster  cremens  368. 
Odontaster  granulosus  367,  372. 
Odontaster  grayi  367,  372. 
Odontaster  hispidus  349. 
Odontaster  pedicellaris  367. 
Odontaster  penicillatus  367,  372. 
Odontaster  pilulatus  367,  372. 
Odontaster  robustus  349. 
Odontasteridae  283,  302. 
Odontophores  37. 

oerstedii,  Astropecten  318,  319,  346,  373- 
Ophiactis  14. 
Ophidiaster  309. 
Ophidiasteridae  282,  285,  308. 
Ophiocoma  pumila,  number  of   rays 

14. 
ornatissimus,   Astropecten   320,   346, 

349- 

ornithopus,  Linckia  310. 
Oreaster  nodosus  286. 
Oreaster  reticulatus  286. 
Oreasteridae  281,  282,  286. 
Orthasterias,  new  genus  48,  64,  168, 

352,  374- 

Orthasterias  biordinata,  new  species 
48,  64,  172,  341 ;  pis.  63,  82. 


402 


INDEX 


Orthasterias  californica,  new  species 

48,  64,  174,  185,  344,  350;  pis.  68, 

70,  80,  81. 
Orthasterias  columbiana,  new  species 

11,  48,  50,  64,  168,  178,  183,  341, 

344,  346,  348;  pis.  24,  35,  65,  78,  79, 

109. 
Orthasterias  dawsoni,  new  species  48, 

65,  172,  175,  342 ;  pis.  23,  75,  80, 

81. 

Orthasterias  eustyla  168. 
Orthasterias  forcipulata  n. 
Orthasterias  forreri  19,  65,  179,  344, 

345 ;  pis.  65,  66,  70,  77,  80. 
Orthasterias    forreri    forcipulata   65, 

1 80,  342;  pis.  62,  70,  88. 
Orthasterias    gonolena,    new    species 

65,  184,  344,  345,  349;  pis.  67,  68, 

69,82. 

Orthasterias  koehleri  65,  175, 183,  342. 
Orthasterias  leptolena,  new  species  66, 

182,  342;  pis.  64,  77. 
Orthasterias  merriami,  new  species  13, 

65,  177,  342  J  pis.  18,  19,  75- 
Orthasterias  (Stylasterias),  new  sub- 
genus  65. 
Orthasterias     (Stylasterias)     forreri 

342. 
Orthasterias  subangulosa,  new  name 

168,  370. 
Orthasterias  tanneri  48,  168,  348,  370; 

pis.  48,  109. 
Ortmannia  205. 
Ossicles,  morphology  of  27. 
Othilia  206. 
(Othilia)    robustus,  Echinaster,  new 

species  207. 

Oyster  beds  damaged  by  starfishes  3. 
Oysters,  food  of  starfishes  3,  5. 
pacificus,  Leptychaster  326,  340,  347; 

pi.  74- 
Psedasterias,  new  genus  352,  353,  355, 

374-  _ 
Paedasterias    chirophora,    new   genus 

355,  371- 

pagenstecheri,  Henricia  363,  372. 
Paleozoic  Echinoderms  12. 
panopla,  Urasterias  52,  182,  370. 
papillosa,  Adelasterias  360,  372. 


papillosa,  Asterias  372. 

papillosa,  Diplasterias  360. 

papillosa,  Ganeria  365,  372. 

papposa,  Asterias  259. 

papposus,   Crossaster  249,   259,   339, 

343 ;  pis.  5,  8,  9,  49. 
papposus,  Solaster  259. 
papposus,  Solaster  (Polyaster)  259. 
papulosus,  Pisaster  n,  58,  68,  91,  340; 

pis.  42,  43,  60,  76,  80. 
Parachasteridce  310. 
Parapatiria  crassa  263. 
Parasterias,  new  genus  53,  55,  66, 187, 

374- 
Parasterias   albertensis,   new   species 

53,  55,  187,  342,  348;  pis.  57,  70. 
Parastropecten  325. 
Parastropecten  inermis  325. 
parvispina,  Evasterias  troschelii,  new 

variety  63,  163,  341. 
Patagonian  species  2,  12,  15,  56,  353. 
patagonica,  Porania  368. 
patagonicus,  Astrogonium  368. 
patagonicus,    Ceramaster    291,    339, 

343,  347,  349,  368. 
patagonicus,  Pentagonaster  291. 
patagonicus,  Pseudar chaster  368. 
Patiria  264. 

Patiria  chilensis  348,  364,  372. 
Patiria  coccinea  263,  373. 
Patiria  gayi  364,  372. 
Patiria  miniata  14,  264,  343,  344,  346, 

347,  348,  3495  Pis.  7,  108,  109. 
Patiria  pectinifera  349. 
Patiriella  372. 

Patiriella  bispinosa  364,  372. 
Patiriella  calcarata  364,  365,  372. 
Patiriella  fimbriata  364,  365,  372. 
Patiriella  pusilla  364,  365,  372. 
Patiriella  regularis  263,  373. 
paucispina,  Asterias  98. 
paucispinus,  Pisaster  19,  98,  340,  344, 

345 ;  pi.  36. 
Paulia  287. 

paxillatus,  Solaster  258,  339,  343,  347. 
Paxillosa,  suborder  24,  283,  313. 
Paxillosa  280,  281. 
Paxillosce  280. 

paxillosus,  Lebrunaster  365,  372. 
Pectinata,  Cribrella  230. 


INDEX 


403 


pectinata,  Henricia  212. 

pectinata,  Henricia  sanguinolenta  229, 

230;  pi.  49. 

pectinifera,  Patiria  349. 
Pectinodiscus  329. 
Pedicellariae  of  Asteriidae  23,  30,  35. 
pedicellaris,  Odontaster  367. 
Pedicellaster  120,  127,  202. 
Pedicellaster  antarcticus  367,  371. 
Pedicellaster  scaber  368. 
Pedicellaster  typicus  202. 
Pedicellasteridae  24,  26,  aoa. 
penicillaris,  Asterinopsis  263,  373. 
penicillatus,  Odontaster  367,  372. 
penicillatus,  Solaster  262. 
Pentaceros  286. 
Pentacerotidce  282,  285. 
pentactis,  Lophaster  368. 
Pentagonaster  286,  288. 
Pentagonaster  289. 

Pentagonaster  austro-granularis  369. 
Pentagonaster  granularis  290. 
Pentagonaster  patagonicus  291. 
Pentagonaster  pulchellus  286. 
Pentagonasterince  283,  285,  289. 
pentagonulus,  Goniodiscus  287. 
peregrinator,  Diplopteraster  368. 
peregrinator,  Retaster  368. 
Peribolaster  biserialis  366. 
Peribolaster  foliculata  366,  372. 
Perknaster  densus  364. 
Perknaster  fuscus  364. 
perrieri,  Anasterias  51,  354. 
perrieri,  Anasterias  355. 
Perrieri,  Asterias  354,  356. 
perrieri,  Asterina  365,  372. 
perrieri,  Asterinides  365,  372. 
perrieri,   Sporasterias  353,  354,  356, 

371- 

Persephonaster  327. 
perspicuus,  Hymenaster  368. 
Pertusa,  Asterias  226. 
pertusa,  Linckia  226. 
Phanerozona,  order  24,  280,  282. 
Phataria  309. 
Phatatria  biserialis  346. 
Phataria  (Linckia)  unifascialis,  var. 

bifcfscialis  309. 
Philomaster  289. 
phragma,  Luidia  367. 


phrygiana,  Hippasteria  286,  301,  371 ; 

pis.  47,  48,  49. 
pilulatus,  Gnathaster  367. 
pilulatus,  Odontaster  367,  372. 
pinguis,  Cycethra  366. 
Pisaster  18,  24,  28,  31,  33,  38,  39,  54, 

56,  67,  72,  352. 
Pisaster  brevispina  77. 
Pisaster  brevispinus  57,  77,  93,  340, 

344,  345;  pis.  41,  44,  45,  69,  76. 
Pisaster  capitatus  19,  57,  81,  344,  345 ; 

pis.  36,  56. 
Pisaster  confertus  57,  73.  167,  340; 

pl.38. 

Pisaster  fissispina  76. 
Pisaster  fissispinus  34,  57,  76,  344 ;  pi. 

39- 
Pisaster  giganteus  n,  57,  84,  87,  89, 

98,  344;  pi.  37. 
Pisaster  grandis  13. 
Pisaster  grayi  58,  91,  97,  344. 
Pisaster  katherina  Per.  58,  89,  91,  97. 
Pisaster  liitkenii  n,  57,  68,  83,  344, 

345,  362. 

Pisaster  liitkenii,  var.  australis,  new 

variety  85,  88,  345- 
Pisaster  liitkenii,  var.  exquiseta  345. 
Pisaster  ochraceus  n,  14,  57,  69,  72, 

340,  344,  345  5  pis.  21,  49,  56. 
Pisaster  ochraceus,  var.  nodiferus  57, 

72,  340,  344  J  pl-  56. 
Pisaster  papulosus  u,  58,  68,  91,  340; 

pis.  42,  43,  60,  76,  80. 
Pisaster  paucispinus  19,  98,  340,  344, 

345  J  pl-  36. 

planci,  Acanthaster,  364,  373. 

planci,  Asterias  364. 

planeta,  Pontaster  369. 

Plates,  skeletal  23,  28,  31. 

plena,  Leptasterias  epichlora,  new  sub- 
species 61,  140,  341. 

Plutonaster  281. 

Podasterias  8,  353,  361. 

Podasterias  loveni  361. 

Podasterias  lutkeni  44,  51,  351,  361, 
372. 

Podasterias  meridionalis  361,  372. 

Podasterias  spinosa  361,  372. 

Podasterias  steineni  351,  361,  372. 

polaris,  Asterias  107,  108,  113,  370. 


404 


INDEX 


polaris,  Ctenodiscus  330. 
polyacantha,  Henricia  213. 
(Polyaster)  papposus,  Solaster  259. 
Polyasterias  45. 

Polyasterias  fernandensis  360,  371. 
polygrammus,  Cosmasterias  360,  371. 
polygrammus,  Stichaster  49,  358,  360, 

polyplax,  Allostichaster  373. 
polyplax,  Asterias  (Stichaster)  125. 
polyplax,  Stichaster  362. 
polythela,  Asterias  59,  104,  338,  347; 

pis.  55,  70,  72,  79,  84. 
Pontaster  planeta  369. 
Pontasterinae  283,  311. 
Porania  281. 

Porania  antarctica  368,  372. 
Porania  glaber  368. 
Porania  magelh&nica  368. 
Porania  patagonica  368. 
Porania  pygmeea  371. 
Poraniidce  304. 
Poraniinae  305. 
Poraniopsis  205. 

Poraniopsis  echinaster  349,  363,  372. 
Poraniopsis  inflata  206,  344,  346,  349, 

363. 

Poraniopsis  inflata  flexibilis  206. 
Poraniopsis  mira  363,  372. 
Porcellanaster  315. 
Porcellanasteridae  283,  328. 
Porcellanasterinae  283,  329. 
praestans,  Cribrella  368. 
praestans,  Henricia  368. 
problema,  Asteracanthion  147. 
procurator,  Ctenodiscus  333. 
Pseudarchaster  285. 
Pseudarchaster  patagonicus  368. 
Pseudarchasterinae  283. 
Pseudoreaster  obtusangulus  287. 
Psilaster  315. 
Psilaster  fleuriaisi  369. 
Pteraster  8,  267. 
Pteraster  affinis  278. 
Pteraster  aporus  277,  278. 
Pteraster  capensis  278. 
Pteraster  carribaeus  278. 
Pteraster  cribrosus  278. 
Pteraster  danae  278. 
Pteraster  gracilis  271,  343,  348. 


Pteraster  hebes  270,  343 ;  pi.  96. 
Pteraster  hexactis  274. 
Pteraster  ingouffi  368. 
Pteraster  lebruni  368. 
Pteraster  marsippus  273,  339,  347. 
Pteraster  militaris  272,  278,  339,  343. 
Pteraster  multipes  278,  279. 
Pteraster  multispinus  271,  343,  348. 
Pteraster  obscurus  13,  17,  274,  339. 
Pteraster  obscurus,  var.  octaster  13, 

276,  339,  347. 
Pteraster  octaster  274. 
Pteraster  pulvillus  273,  278,  339. 
Pteraster  reticulatus  268. 
Pteraster  rugatus  278,  368. 
Pteraster  semireticulatus  278. 
Pteraster  stellifer  278,  368. 
Pteraster  (Temnaster)   hexactis  274. 
Pteraster  tesselatus  268,  271,  339,  343, 

348;  pis.  32,86,  97. 
Pteraster  tesselatus  arcuatus  269. 
Pteraster  tesselatus  hebes  270. 
Pterasteridae  204,  266. 
Pterasterides  277. 
Pterasterides  aporus  278,  339,  347. 
Pterasterinae  266,  267. 
pugetana,  Leptasterias  epichlora,  new 

subspecies  61,  142,  341. 
pulchellus,  Pentagonaster  286. 
pulvillus,  Pteraster  273,  278,  339. 
pusilla,  Patiriella  364,  365,  372. 
Pycnopodia  4,  6,  12,  15,  39,  40,  43,  67, 

68,  197- 
Pycnopodia  helianthoides  n,  35,  198, 

342,  344,  348;  pis.  29,  30,  31,  73, 

74,88. 

Pycnopodida  197. 
Pycnopodiinae  26,  197. 
pygmaea,  Asterina  371. 
pygnuea,  Porania  371. 
pygmeeus,  Ctenodiscus  371. 
Pythonaster  267. 
Pythonasteridae  204. 

radiosus,  Labidiaster  352,  371. 

Randasia  287. 

Rathbun,  Richard,   species  dedicated 

to  190. 

Rathbunaster  12. 
Rathbunaster  californicus  197. 


INDEX 


405 


rathbuni,   Allasterias   53,   55,   66,   67, 

189.  337,  338,  347- 
rathbuni  nortonensis,  Allasterias  67, 

191.338,  347;  pi.  78. 

Rays,  development  28. 

Rays,  number  and  variability   12-15, 

38. 

Rays,  spontaneous  fission  39. 
regularis}  Cycethra  366. 
regularis,  Leptasterias  epichlora,  new 

variety  61,  139,  341. 
regularis,  Patiriella  263,  373. 
regularis,  Solaster  368. 
Reproductive  organs  32. 
Retaster  gibber  366,  372. 
Retaster  multipes  279. 
Retaster  peregrinator  368. 
Retaster  verrucosus  366,  368,  372. 
reticulatus,  Gonaster  286. 
reticulatus,  Oreaster  286. 
reticulatus,  Pteraster  268. 
Rhopia  209. 
Rhopia  sepositus  363. 
ritteri,  Bunodaster  316,  322,  345,  348; 

pis.  86,  104,  105. 
robusta,  Ganeria  365. 
robusta,  Stolasterias  373. 
robusta,  Stylasterias  350,  373. 
robustus,  Echinaster  342. 
robustus,  Echinaster   (Othilia),  new 

species  207. 

robustus,  Odontaster  349. 
rosea,  Asterias  40,  362. 
rosea,  Cribella  226. 
rosea,  Stichastrella  40,  370. 
rosea,  Stichaster  40. 
rubens,  Asteracanthion  373. 
rubens,  Asterias  44,  45,  68,  188,  337, 

346. 

rubens,  Asterias  107,  189. 
rudis,  Evasterias  troschelii,  new  vari- 
ety 63,  158,  34L 
rudis,    Henricia    sanguinolenta,    new 

variety  233,  338,  347 ;  pi.  86. 
rugatus,  Pteraster  278,  368. 
rugispina,  Asterias  51,  53,  353,  355. 
rugispina,  Sporasterias  353,  356,  357, 

371- 
rupicola,  Asterias  356. 


rupicola,  Sporasterias  51,  53,  353,  356, 

371. 
rustica,  Asterias  357. 

saanichensis,  Asterias  132,  135. 
sanguinolenta,  Asterias  226. 
sanguinolenta,  Cribrella  223,  226. 
sanguinolenta,  Echinaster  226. 
sanguinolenta,   Henricia   n,   14,  212, 

226,  338,  342 ;  pi.  49,  88. 
sanguinolenta  miliaris,  Henricia,  new 

subspecies  234,  370;  pi.  88. 
sanguinolenta,  var.  pectinata,  Henricia 

338;  pi.  49. 
sanguinolenta,   var.   rudis,   Henricia, 

new  variety  233,  338,  347;  pi.  86. 
sarsii,  Echinaster  226. 
scaber,  Pedicellaster  368. 
scalprifera,  Smilasterias  53,  371. 
Sclerasterias  guernei  44,  50,  370. 
scrobiculata,  Echinaster  227. 
sebce,  Goniodiscides  288. 
semireticulatus,  Pteraster  278. 
Senses,  instincts,  memory  9. 
sentus,  Echinaster  207. 
sepositus,  Rhopia  363. 
sertulifera,  Asterias  100. 
sertulifera,  Marthasterias  19,  58,  100, 

345,  349- 

sexradiata,  Henricia  371. 
siderea,  Enoplopatiria  365,  373 ;  pi.  109. 
siderea,  Leptasterias  epichlora  alas- 

kensis,  new  variety  61, 137,  341. 
siderealis,   Astropecten,   new   species 

317.  345,  346,  349J  pl-  50- 
simplex,  Cribrella  368. 
simplex,  Cycethra  366,  372. 
simplex,  Henricia  368. 
singularis,  Asterodon  367,  372. 
Skeletal  plates,  growth  of  31. 
Skeleton  of  starfishes  20. 
sladeni,  Cribraster  363,  372. 
Smell,  sense  of  9. 
Smilasterias  44,  53. 
Smilasterias  scalprifera  53,  371. 
smithii,  Echinaster  368. 
Solaris,  Acanthaster  364,  372. 
Solaris,  Echinaster  364. 
Solaster  4,  13,  15,  17,  18,  242. 
Solaster  australis  366,  368,  372. 


406 


INDEX 


Solaster  constellatus  257,  343,  348; 

pis.  46,  90,  93,  94. 
Solaster  dawsoni  349,  339,  342,  348; 

pis.  46,  90,  91,  92. 

Solaster   dawsoni   arctica,   new   sub- 
species 252,  253,  347;  pi.  87. 
Solaster  decemradiatus  254. 
Solaster  endeca  n,  243,  344,  248,  249, 

339;  pis.  9,87,  89. 
Solaster  galaxides  248,  342,  348;  pis. 

46,  87,  89. 

Solaster  octoradiatus  368. 
Solaster  papposus  259. 
Solaster  paxillatus  258,  339,  343,  347 
Solaster  penicillatus  262. 
Solaster  (Polyaster)  papposus  259. 
Solaster  regularis  368. 
Solaster  stimpsoni  250,  254,  339,  348; 

pis.  10,  II,  15,  46,  86,  94,  95. 
Solaster  subarcuatus  368. 
Solaster  vancouverensis  254. 
Solasteridae  41,  204,  242. 
Solasterince  242. 
spatulifera,  Henricia  224. 
spatulifera,  Henricia  leviuscula,  var. 

224,  342,  344,  346 ;  pis.  5,  14. 
Species,  determination  of  17. 
spectabile,  Asteracanthion  358. 
spiculifera,  Cribrella  220. 
spiculifera,  Henricia  220. 
spiculifera,  Henricia  leviuscula  220, 

338, 342 ;  pis.  87, 107. 
Spines,  34,  36,  38. 
spinosa,  Diplasterias  361,  372. 
spinosa,    Hippasteria   301,   339,   343, 

345,346,  348;  pis.  50,98. 
spinosa,  Podasterias  361,  372. 
spinosus,  Echinaster  207. 
Spinulosa  24,  203. 
spirabilis,  Asterias  51,  354. 
spirabilis,  Sporasterias  354. 
spitzbergensis,  Asterias  148,  150. 
spitzbergensis,    Ctenasterias    53,    55, 

148,  370. 

Sporasterias  8,  53,  350,  353,  355. 
Sporasterias   antarctica   51,   53,   353, 

354,  371. 

Sporasterias  cocosana  350. 
Sporasterias  cunninghami  353. 
Sporasterias  galapagensis  350. 


Sporasterias  hyadesi  354. 
Sporasterias  mariana  350. 
Sporasterias  perrieri   353,  354   356, 

37i. 
Sporasterias  rugispina  353,  356,  357, 

371. 
Sporasterias  rupicola  51,  53,  353,  356, 

37i. 

Sporasterias  spirabilis  354. 
Sporasterias  varia  353. 
Sports  or  hybrids  17. 
steineni,  Asterias  361. 
steineni,  Diplasterias  361. 
steineni,  Podasterias  351,  361,  372. 
stellans,  Lophaster  366,  368,  372. 
stellatus,  Mediaster  298. 
Stelleridae  f  orcipulatae  24. 
Stelleridae  Spinulosae  203. 
stellifer,  Pteraster  278,  368. 
stellionura,  Asterias  51. 
Stellonia  101,  206. 
Stenasterias,  new  genus  145,  374. 
Stenasterias  macropora  13,  66,   145, 

341,  348;  pis.  50,  74,  84. 
Stephanaster  286. 
Stephanasterias  36,  55,  130,  146. 
Stephanasterias  albula  39,  40,  41,  55, 

6>,  147,  338. 

Stephanasterias  albula,  var.  nitida  147. 
Stephanasterias  gracilis  147. 
stichantha,  Distolasterias  44,  49,  185, 

373- 

stichantha,  Stolasterias  47. 
Stichaster  30. 
Stichaster  146. 
Stichaster  albulus  41,  147. 
Stichaster  aurantiacus  40,  362. 
Stichaster  f  elipes  49. 
Stichaster  nutrix  51,  351. 
Stichaster  polygrammus  49,  358,  360, 

371. 

Stichaster  polyplax  362. 
Stichaster  rosea  40. 
Stichaster  striatus  40,  362,  372. 
Stichasteridae  27,  42. 
Stichasterinae  26,  39,  40,  42,  146. 
Stichastrella,  new  genus  40,  374. 
Stichastrella  rosea  40,  370. 
stimpsoni,  Solaster  250,  254,  339,  348; 

pis.  10,  ii,  15,  46,  86,  94,  95. 


INDEX 


407 


Stolasterias  43,  44,  46,  350. 
Stolasterias  45,  47- 
Stolasterias  alexandri  372. 
Stolasterias  candicans  367. 
Stolasterias  neglecta  47- 
Stolasterias  robusta  372. 
Stolasterias  stichantha  47. 
(Stolasterias)    tenuispina,    Coscinas- 

terias  46,  49. 

Stolasterias  volsellata  46. 
stolidota,    Calvasterias   42,   354,    357, 

371. 

Stomach  of  starfishes  21. 
striattis,  Stichaster  40,  362,  372. 
studeri,  Anasterias  355,  371. 
studeri,  Asterias  51. 
studeri,  Gastraster  360,  372. 
studeri,  Henricia  363. 
Stylasterias,  new  subgenus  33,  48,  65, 

179.  374- 

Stylasterias  forreri  48,  50. 
Stylasterias  neglecta  48,  370. 
Stylasterias  robusta  350,  373. 
subangulosa,  Orthasterias,  new  name 

168,  370. 

subarcuatus,  Solaster  368. 
subelectilis,  Cycethra  366. 
subnodosa,  Evasterias  troschelii,  new 

variety  63,  163,  341. 
subnodulosa,    Leptasterias    epichlora, 

new  variety  62,  139,  341. 
subregularis,    Leptasterias    epichlora, 

new  variety  62,  73,  139,  341. 
Subspecies,  definition  of  17. 
Sucker-feet  4,  6,  15,  39. 
sulcifera,  Asterias  48,  358,  361. 
sulcifera,  Cosmasterias  48,  358,  360, 

361,  371- 
tanneri,  Orthasterias  48,  168,  348,  370 ; 

pis.  48,  109. 
Temnaster  267. 
Temnaster  hexactis  267,  274. 
Temperature  during  Glacial  Period  2. 
Temperature  of  coast  waters  2,  3. 
Temperature  of   water  limits  distri- 
bution 19. 

tenera,  Anasterias  355,  371. 
tenera,  Leptasterias  8,  37O. 
tenuispina,  Coscinasterias  38,  43,  45, 

370. 


tenuispina,  Coscinasterias  (Stolaster- 
ias) 46,  49. 

tenuispinus,  Echinaster  208,  346,  349, 
373 ;  pi.  107. 

tesselatus  arcuatus,  Pteraster  269. 

tesselatus,  Goniaster  286. 

tesselatus  hebes,  Pteraster  270. 

tesselatus,  Pteraster  268,  271,  339, 
343,  348;  pis.  32,  86,  97- 

Thyone  briareus  10. 

tomidata,  Asterias  (Cosmasterias)  48. 

tomidata,   Cosmasterias  48,  50,  359, 

371. 

Tonia  atlantica  40,  362. 

Tosia  287,  288,  289. 

Tosia  arctica  292. 

Tosia  (Ceramaster)  granularis  290. 

Tosiaster,  new  genus  292. 

Tosiaster  arcticus  292,  339;  pis.  50, 

99. 

Touch,  sense  9. 
Tremaster  mirabilis  371. 
trosfhelii,  Asterias  151. 
troschelii,  Evasterias  n,  36,  51,  54,  63, 

114,  151,  156,  341,  344,  348;  pis. 

22,  25,  26,   I06. 

troschelii,  var.  alveolata,  Evasterias, 
new  variety  63,  157,  162,  341 ;  pi. 
62. 

troschelii,  var.  densa,  Evasterias,  new 
variety  63,  161,  341. 

troschelii,  var.  parvispina,  Evasterias, 
new  variety  63,  163,  341. 

troschelii,  var.  rudis,  Evasterias,  new 
variety  63,  158,  341. 

troschelii,  var.  subnodosa,  Evasterias, 
new  variety  63,  163,  341. 

tumida,  Henricia  214,  334,  239,  240, 
338,  347;  pis.  12,87. 

tumida  borealis,  Henricia,  new  sub- 
species 236,  339,  342,  347 ;  pis.  12, 
88. 

turqueti,  Asterias  372. 

turqueti,  Cryptasterias  362,  372. 

turqueti,  Diplasterias  362. 

Tylaster  305. 

typicus,  Pedicellaster  202. 

typus,  Linckia  211. 


408 


INDEX 


unifascialis,  var.  bifascialis,  Phataria 

(Linckia)  309. 
Uraster  101. 
Urasterias  33,  34,  5*.  55,  66,  187,  352, 

374- 

(Urasterias)  forcipulata,  A sterias  180. 
Urasterias  gunneri  52. 
Urasterias  linckii  33,  51,  55,  66,  181, 

187,  370;  pi.  70. 
Urasterias  panopla  52,  182,  370. 

Valvaster  283. 

Valvasteridae  282. 

Valvasterina  283. 

Valvata,  order  280,  282,  284. 

Valvatce  280. 

Valvulatce,  order  284. 

valvulifera,    Dermasterias    imbricata 

305,  308,  343. 

Valvulosa,  suborder  24,  282,  284. 
vancouverensis,  Solaster  254. 


Vancouver*,  Asterias  13,  125. 
vancouveri,  Leptasterias  61,  125,  340. 
varia,  Asterias  51. 
varia,  Sporasterias  353. 
Varieties,  definition  17. 
variolata,  Linckia  211. 
Velata,  suborder  204,  266. 
verrillii,  Anasterias  354,  355,  37 1- 
verrucosa,  Cycethra  366,  372. 
verrucosus,  Diplopteraster  368. 
verrucosus,  Retaster  366,  372. 
versicolor,  Allasterias  106,  373. 
victoriana,  Asterias  59,  102,  340,  348; 

pi.  53,  54,  69,  82. 
violacea,  Asterias  107. 
volsellata,  Heterasterias  27, 47, 49, 373. 
volsellata,  Stolasterias  46. 
vulgaris,  Asterias  38,  370. 

Zoroasteridae  24,  26,  41. 


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